VOLUME 58, NUMBER 3 JULY-SEPTEMBER 2011 RECONSIDERATION OF THE TAXONOMIC STATUS OF MASON’S LILAEOPSIS — A STATE-PROTECTED RARE SPECIES IN CALIFORNIA Peggy L. Fiedler, Esa K. Crumb, and A. Kate Kn0Xx..........ccccccceeeeeeeeeereees 131] A COMPARISON OF THE EFFECTS OF NA2SOq4 AND NACL ON THE GROWTH OF HELIANTHUS PARADOXUS AND HELIANTHUS ANNUUS (ASTERACEAE) MoO. Mendez and O; W, Van AUbeH sivtecc Met accsesenstoomee eee tona eater ondees 145 THE DIVERSITY AND BIOGEOGRAPHY OF THE ALPINE FLORA OF THE SIERRA NEVADA, CALIFORNIA SITU TTD WV RUNG CP os ese ch ace nee eA RD wk ee RM PR Anan ounesedausewadeass 53 A NEw SPECIES OF ASTRAGALUS (FABACEAE) FROM THE WASATCH MOUNTAINS OF UTAH DCU TCOWE: COP DIM oh cients aiecdet as ORR See AON hens oli Sulla cbueaeenaule 185 GRIMMIA VAGINULATA, (BRYOPSIDA, GRIMMIACEAE) A NEW SPECIES FROM THE CENTRAL COAST OF CALIFORNIA TR CATLCLIA INCU VIIIOTA te iN rene Le SN eh 190 INTRODUCTION TO CALIFORNIA CHAPARRAL CGHAESTODICTAIBUGV LOCK. cisirs caissad Mccre tetec ae Oe 91g siRNA ned as Fes ees wae snide 199 COP ANTEINEE GOIRINITIN reteset ite cia sts ano, aeRO Me cists cscs a wits Roe oak oes Sanne vamaeemniads 201 DEIN eee se Bites aah ee iste casa ca camnem sO taneaaenaieasoadeenuasiheeeueneliel 204 IN TERS (610 SAR te oe) ee ere eee D205 ManbroNo (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. 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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: Marc Los Huertos, Division of Science & Environmental Policy, California State University, Monterey Bay, Seaside, CA 93955, mloshuertos@csumb.edu Recording Secretary: Mike Vasey, Department of Biology, San Francisco State University, San Francisco, CA 94132, mvasey @sfsu.edu Corresponding Secretary: Heather Driscoll, University Herbarium, University of California, Berkeley, CA 94720, hdriscoll @ berkeley.edu 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, 3060 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: Ben Carter, Department of Integrative Biology and University Herbarium, University of California, Berkeley, CA 94720, bcarter@berkeley.edu. Webmaster: Susan Bainbridge, Jepson Herbarium, University of California, Berkeley, CA 94720-2465, sjbainbridge @ berkeley.edu. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). MADRONO, Vol. 58, No. 3, pp. 131-144, 2011 | GMT HSONIay APR 129012 LIBRARIES RECONSIDERATION OF THE TAXONOMIC STATUS OF MASON’S LILAEOPSIS — A STATE-PROTECTED RARE SPECIES IN CALIFORNIA PEGGY L. FIEDLER Natural Reserve System, UC Office of the President, Office of Research and Graduate Studies, 1111 Franklin Street, 6" Floor, Oakland, CA 94607 peggy.fiedler@ucop.edu ESA K. CRUMB Wetlands and Water Resources, 818 Fifth Ave, STE 208, San Rafael, CA 94901 A. KATE KNOX 7730 19% Avenue NE, Seattle, WA 98115 ABSTRACT Lilaeopsis masonii is a California state-listed rare species with a wide range of morphologies observed in the field throughout its range, and in herbaria collections. This extensive variation confounds reliable taxonomic identification, particularly for those specimens intermediate between L. masonii and its sister taxon, L. occidentalis. To investigate the genetic basis of this morphological variation, we examined two portions of the Lilaeopsis genome in seven species. Specifically we sought to determine whether L. masonii is sufficiently distinct from its closely related, widespread congener to continue to warrant specific status. DNA sequence analysis of ITS1, 5.8S, and ITS2 nuclear ribosomal DNA revealed no differences between L. occidentalis and L. masonii California collections, and minimal differences between these samples and L. occidentalis collected from the state of Washington, suggesting strongly that these two species form a single clade. A combination of fragment data from three AFLP primers yielded 274 fragments from 29 samples. Genetic Manhattan distance values calculated from the AFLP matrix within species ranged from a low of 1.4 to a high of 6.6, reflecting minor differences among all samples. UPGMA cluster phenograms support the results of the PCA analysis, illustrating a cluster of L. occidentalis + masonii samples distinct from other Li/aeopsis species. Because conservation dollars should protect unique evolutionary entities, we suggest that L. masonii be subsumed under L. occidentalis and therefore no longer receive formal state protection. Key Words: AFLP, Apiaceae, California endangered species Act, goldilocks conundrum, ITS, lilaeopsis masonii, lilaeopsis occidentalis, UPGMA. Lilaeopsis masonii Mathias & Constance (Ma- son’s lilaeopsis) is one of 15 wetland or aquatic species of the widespread genus Lilaeopsis Greene within the Apiaceae. The genus Lilaeopsis 1s comprised of perennial herbs characterized by a horizontal stem with leaves commonly in clusters (“‘ramets’’) borne directly from the stem, although rarely leaves occur individually. Lilaeopsis is notable in its morphologic simplicity—entire, generally linear leaves; simple umbels; absence of a carpophore; and, a strongly reduced habit (Petersen et al. 2002; Downie et al. 2000; Downie et al. 2008). Such simple morphology has led to a long history of taxonomic uncertainties and difficulty in the reconstruction of its phylogeny. Evidence for monophyly of Li/aeopsis is strong (Petersen et al. 2002). However, recent research based on molecular evidence from nuclear and chloroplast genes suggests that the genus is best placed in the Oenantheae tribe within the Apioideae (Downie et al. 2008) and that Lilaeop- sis 18 sister to the clade comprising Ptilimnium, Limnosciadium, Daucosma, Cynosciadium and rachis-leaved species of Oxypolis, not the Mexi- can genus Neogoezia as suggested by Petersen et al. (2002). The New World endemics clade of tribe Oenantheae is native to North America and comprises a monophyletic group that appears to be evolving much faster than any other major clade recognized in the tribe (Hardway et al. 2004). Early taxonomic work on the genus in California by Hill (1927) and Mason (1957) included mention of comparatively smaller and narrower leaves in Lilaeopsis specimens occurring away from the coast, in contrast to a relatively more robust coastal form. Professor Herbert Mason, an early expert on the wetland flora of California, first collected a relatively smaller Lilaeopsis from Brannan Island of the San Francisco Bay/Sacramento-San Joaquin Delta (Bay-Delta). He referred to the smaller form as the “San Francisco Bay and river-mouth” form (Mason 1957: 631). This specimen, according to Mason (unpublished) was ‘‘definitely distinct from the coastal L. occidentalis.’’ Western lilaeopsis (Lilaeopsis occidentalis J. M. Coulter & Rose) is a widespread, common species, 139 MADRONO ranging from the Queen Charlotte Islands of British Columbia, Canada to Marin County, California (Affolter 1985). Considered to be a coastal species confined to salt water or brackish water intertidal habitats, collections of L. occi- dentalis from inland fresh water lentic and lotic habitats are known, but considered “‘uncharac- teristic’ (Affolter 1985). Lilaeopsis masonii was not described as a distinct taxon for two decades after the smaller form in the Bay-Delta was first observed. In 1977, Mathias and Constance formally recog- nized the diminutive nature of a specimen obtained from Twitchell Island in the Bay-Delta as L. masonii (Mathias and Constance 1977). Mathias and Constance described L. masonii as distinct from L. occidentalis based upon the former (rare) taxon bearing narrower, typically shorter, and more or less terete leaves, and an inland distribution. They honored Herbert Ma- son’s expertise in the wetland flora of the State with the specific epithet. Mason’s lilaeopsis was one of the first vascular plant species to be protected as “‘rare’”’ under the California Endangered Species Act (CESA) (California Fish & Game Code 882050, ef seq.). At the time of its listing in November 1979, only seven population occurrences were known (CNDDB 2009). Since formal protection, the documented extent of geographic distribution and population abundance of L. masonii has increased nearly three-fold, primarily as a result of concentrated field survey efforts conducted in the early 1990’s by Golden, Fiedler, and Zebell (Golden and Fiedler 1991; Golden 1992; Fiedler and Zebell 1993; Zebell and Fiedler 1996). Today, Mason’s lilaeopsis is known to occur within 24 USGS quadrangles and seven counties (CNPS 2008), spanning across roughly 690 square miles. One hundred eighty-six documented occurrences are on record with the state (CNDDB 2009), although most, but not all are extant. A History of Taxonomic Uncertainty Confusion over the taxonomic limits of this rare species existed from the beginning of its description. Two examples are relevant. First, a long-controversial Lilaeopsis specimen collected by Schreiber (#42266 UC, 28 June 1936) from Chicken Ranch Beach in Marin County derives from outside the circumscribed geographic range of the endemic inland taxon. Leaf lengths from this specimen range between 15 to 42 mm, a morphological range characteristic of compara- tively larger leaf lengths for L. masonii. However, it is possible to key these larger leaved Chicken Ranch Beach specimens to L. occidentalis in every relevant flora (e.g., Hickman 1993). Affolter (1985) examined this specimen in his monograph of Lilaeopsis, and accepted it as L. [Vol. 58 masonii, but noted that it was a geographical outlier for the rare, Bay-Delta endemic species. Today, CNPS (Tibor 2001) acknowledges that this specimen is likely to be L. occidentalis, not the rare L. masonii, but provides no explanation. Several attempts to relocate this Lilaeopsis material at Chicken Ranch Beach by the authors have failed as the population appears to be extirpated, thereby making an independent spe- cies corroboration impossible. It is important to note that (1) numerous collections of L. occidentalis from the beaches of Marin and Sonoma counties exist, (2) leaf lengths range by an order of magnitude or more within and between adjacent populations of L. occiden- talis, (3) the number and clarity of internal crosswalls considered important diagnostic char- acters are more likely a function of relative plant size, exposure, or both, and (4) inland collections of the common species are known from the state of Washington (e.g., UC 1594452; 4 September 1962). Also noteworthy, L. masonii has never again been collected on the Pacific coast of North America beyond Schreiber’s Marin Co. collection in 1936. Further, Affolter (1985) remarked that leaves from a collection of L. masonii (derived from Sherman Island immediately down river of Twitchell and Brannan islands) cultivated for his greenhouse comparisons were “‘remarkably longer than any of the herbarium material” (Affolter 1985:70). He suggested the observed overall larger and more robust greenhouse material was evidence of how difficult it is to understand vegetative plasticity from herbarium material alone. However, the relatively robust response of Mason’s lilaeopsis to the mild conditions of a greenhouse suggests strongly that strict morphological distinctions between the two taxa are problematic. Additional morphological characters further support the assertion that L. masonii is not distinctly different from L. occidentalis. Affolter (1985:70) noted that the “two taxa are similar in several respects,”’ including similar (1) leaf shapes (linear), (2) rhizome branching architecture, (3) fruit shapes, (4) fruit cell types, (5) fruit venation patterns, (6) habitats, and they have (7) overlap- ping geographic distributions. Despite all these similarities, Affolter (1985:71) supported their separate specific status, primarily because ““when grown under a common-garden environment in the greenhouse, the two species retained the vegetative characteristics that distinguish them in the field.” Subsequent laboratory studies conducted by the principal author and her students (Golden and Fiedler 1991; Golden 1992; Fiedler and Zebell 1993; Zebell and Fiedler 1996) have provided little clarity. Most importantly, no nucleotide variation was found among nine 2011] populations of L. masonii or between L. occiden- talis and L. masonii when the 204 nucleotides of the ITS2 nuclear genome were ascertained (Fiedler and Zebell 1993). Fiedler and her colleagues thus concluded tentatively that the rare species was most likely an inland ecotype not clearly distinct from its widespread congener. Field Observations Decades of field observations of Lilaeopsis throughout the Bay-Delta, Suisun Marsh, and Napa River ecosystems do not reinforce many of the conclusions offered by Hill (1927), Mason (1957, unpublished), Mathias and Constance (1977), and Affolter (1985) supporting the recognition of two distinct taxa. Rather, the few vegetative characteristics that typify this genus are highly variable both within and between populations throughout this region. Occurrences of L. masonii in the lower Napa River, studied since 2001 (WSP 2007, unpublished; Blasland, Bouck, & L, Inc. unpublished; Entrix unpub- lished; L.C. Lee & Associates unpublished; Still- water Sciences and Fiedler unpublished) include a full spectrum of individual ramet sizes. Often, both large and small forms of Lilaeopsis species, easily identifiable to the two different species, can be found growing in the same location. Often the plant stature/leaf length size gradient runs per- pendicular to the shoreline, where the small ‘“masonii’ form (approx.1.5—4.5 cm in height) grows relatively close to the water’s edge, while increasing larger and more robust “‘occidentalis”’ (approx. =11 cm in height) can be found further from the water. “Intermediate” or medium-sized Lilaeopsis material (approx. >6.25 and <11 cm in height) is common throughout this shoreline/ river bank habitat and geographic range, and keys to either (or both) the rare or the common species. We call this phenomenon—.e., range in size of a critical morphological character, with significant overlap between taxa—the ‘‘Goldi- locks Conundrum”’ to highlight the problem that the intermediate-sized material is not “‘just right,”’ but rather, highly problematic. To resolve our conundrum and determine whether L. masonii is a discrete species distinct from L. occidentalis, we initiated a genetic analysis of seven species of this genus. We hypothesized that there were no significant differences between diagnostic portions of the genome selected for this study of the two species, L. masonii and L. occidentalis. Based on these analyses, we then explored whether L. masonii watrants continued recognition as a distinct species or rather, should be subsumed under the widespread and common L. occidentalis. If no significant differences were shown to exist be- tween diagnostic portions of the L. occidentalis and L. masonii genomes, then L. masonii should FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS 133 be subsumed within L. occidentalis, and contin- ued protection under the California Endanger- ed Species Act for L. masonii should be reconsidered. Fallon (2007) noted that genetic information is being used increasingly to resolve taxonomic issues for protection at the federal level under the U.S. Endangered Species Act of 1973 (ESA). She conducted a review of listing decisions made by the U.S. Fish & Wildlife Service and the National Marine Fisheries of species, subspecies, or distinct population segments (DPSs) proposed for protection under the ESA. Fallon determined that the listing fate of a DPS based upon data from more than one genetic marker resulted in a higher probability of protection than candidate taxon or population segment whose discreteness was determined by a single genetic marker. With the cautionary tale of Fallon’s findings in mind, we examined the ITS region of the nuclear genome and, to corroborate our ITS findings, conducted an amplified fragment length poly- morphism (AFLP) analysis on a similar suite of taxa. We chose the ITS region in large part because Hardway et al. (2004) found evidence for particularly rapid evolution in the Oenanthe clade that includes Lilaeopsis when compared to the rest of the taxa. Sequence divergences in this clade averaged 6—7 times higher (approx. 17%) than between species in Oenanthe (approx. 2.8%) or Cicuta (approx. 2.4%) (Hardway et al. 2004). AFLP analysis was selected as a secondary marker system based on the increasing popularity of this form of DNA fingerprinting as a complementary system in phylogenetic studies (Holland et al. 2008). Additionally, AFLP fingerprinting offers a reliable, robust, and genomically comprehensive method of genetic analysis for taxa lacking complex nuclear and organellar markers (Vos et al. 1995). MATERIALS AND METHODS Field Collection Lilaeopsis masonii specimens were collected in the spring of 2007 from locations along the Napa River and in the Sacramento/San Joaquin Delta. Lilaeopsis occidentalis was collected from Bodega Head, California, and Mason and Lawrence lakes in Washington State (Fig. 1). Leaf material to be used in DNA extraction was preserved in silica gel at the time of collection. Vouchers were deposited at the herbarium at San Francisco State Univer- sity (SFSU) (Table 1). Material for L. brasiliensis (Glaz.) Affolter and L. mauritiana G. Petersen & J. Affolter was obtained from a commercial aquarium supplier (freshwateraquariumplants.com). The dataset is composed of 35 nrDNA ITS sequences representing seven taxa, including three sequences MADRONO [Vol. 58 Washington Samples hal ell Br | Maxw ~sa Nevada ‘Hill Slough/Grizzly Island Liberty Island [Twitchell Island 134 = i 7 California 7 Napa’ Bodega “Vv ¢ Collinsville 3 Mile Slough Brannan Is. Boat Ramp — Fic. 1. this study. from Genbank (L. carolinensis J. M. Coulter & Rose, L. novae-zelandiae (Gand.) A. W. Hill, and L. occidentalis), two specimens from the aquarium trade labeled as L. brasiliensis and L. mauritiana, and two specimens of L. schaffneriana (Schltdl.) J. M. Coulter & Rose subsp. recurva (A. W. Hill) Affolter courtesy of the Desert Botanical Garden staff. Comprehensive sampling was conducted for L. masonii and L. occidentalis as the purpose of this study was to resolve the taxonomic classifi- cation for these two species. A detailed systematic O \3sr5 75 150 Miles Map of the geographic locations of Lilaeopsis masonii and L. occidentalis specimens collected for study for Lilaeopsis is in preparation (S. Downie, Univ. of Dlinois, Urbana-Champaign, personal communication). ITS Methods DNA from leaf tissue of five of the seven species was extracted using the Qiagen DNeasy Plant Mini Kit (QIAGEN, Inc., Valencia, CA), following the manufacturer’s protocol with slight modifications. Lilaeopsis carolinensis and _ L. 135 MSAS NSAS OLSAS OSAS ‘OZ*D se[snog ‘U0sIIO “VSN OSHS OSHS OSHS UdsI3}og IID ‘yJNO ‘puevles7 MONT OSAS IOyONOA ON OSAS OSAS OSAS OSHS OSHS OSAS OSAS OSAS OSAS OSHS OSAS OSAS OSAS OSAS OWSAS OSAS OLSAS OSAS OSAS OWSAS OSAS suspley s1urj0g YSN JO “AtUs) ‘yn ‘euNnUasIV FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS UOTBOOT JBYONOA =psz9][09 318Q] ONTHLYON 2011] V/N V/N CO6I/LI/9 1661/7 1/9 V/N 800C/S C/L 800C/S C/L LOOC/L/OI V/N LOOC/LC/8 LOOC/9C/8 LOOC/tC/8 LOOC/VC/8 LOOC/VC/8 LOOC/€C/8 LOOC/VC/8 LOOC/t 1/6 LO0C/9/¢ LOOC/VC/V LOOC/VC/V LOOC/V C/V LOOC/V C/V LOOC/S C/V LOOC/S C/V LOOC/VC/v LOOC/OI/S LOOC/8/S LOOC/8/S LOOC/6/S LOOC/6/S LOOC/8/S LOOC/9/S VN V/N V/N O69TOSE ‘xoiddy Te96LPe ‘xoiddy V/N tVPL8sIs DLOIPCS Oct lecr V/N vILvlcr Ocelecr OO6LECL 8Pe9Icr Ocrél cr 8CS8ICV 9V80ICS S686ECL L8CVECY 6tLLecVy 6CLLecV 99OLLECY 99OLLECV C8S8ECP 86C8ECV OLOSECL O9CLECYL C8C9CCH C8C9CCH LOL9tCcY StO9ECL VCOVECL VCOVECL VN V/N V/N V869CS ‘xoiddy 600LSS ‘xolddy V/N LSecOSo LStcOso 9C8P6r0 V/N 6¢60090 68SS8S0 6vS¢190 SSPst90 SL6rv190 tc6c190 8IL16SO tI Lc9s SeSc9so c9Vc9SO c9Vc9SO0 T19c9SO LT9c9SO0 888c9S0 CS8C9SO 6¢8c9SO OISc9SO 6LEc9SO 6LEc9SO cOEc9SO 8LECc9SO 69¢c9S0 69¢c9S0 VN ONILSV4A VN VN V/N V/N V/N LO! LO! STI V/N STI SIT SII SII SII STI SII SII STI SII SII STI STI SII STI SII STI SII SII SII SIT SII SII VN WL V/N V/N ZV ‘eyOuog ZV ‘astyoog V/N VM ‘UO}sINY ], VM ‘uose|] VO ‘UHR V/N VO ‘ourjog VO ‘ourfos VO ‘ourjos VO ‘olUSTTReIDeS VO ‘olustTRIDeS VO— So yUSTTRIDeS V°— ‘B1SOD PIJUOD Vo ‘eden Vo ‘eden Vo ‘eden Vo ‘eden Vo ‘eden Vo ‘eden Vo ‘eden Vo ‘eden VO ‘eden Va ‘eden Vo ‘eden VO ‘eden Vo ‘eden Vo ‘eden Vo— ‘eden Vo ‘eden VN 97e1s ‘AyUNOD €€TLPIOH F#// Woosyuefdumnuenbesayemysasy PETLPIOH #// Wuoo'sjuv;dumuenbessyemyso1f TETLPIOH F#// PUOZITY “UOkURD BODES I€TLPOOHF #// PUOZITY “UOCKURD BODES CVCOIEAV SLI CPTLPIOH #// VM ‘OOUDIMET YLT precTLP9OH #// VM ‘AAeT Uosepy 9€TLPYOHF // PRAH eSapog 8LC99VAV-SLI OPZLP9OH #// LOO-AIASUT[OD THTLPIOH #// 900-PURTST ATZZIID/Yysnojs [tH SETLPIOH// SOO-PURIS] A9qQrT OOLPEIOH# / /POO- PURIST [24M L IpCLpPOOH# #// €00- dures yeoq purysy uUeuURIg S~TLPIOH #// TOO- UBNOTS AHA € LETLPYOH# //100- JOqIeH AoayoW 6ETLPIOH// LO-ME - Jeary eden ZOH/TOO-VI JoaTy eden Q7OE-MZT JoaTy eden B7OE-MC Joary eden QLOA-AT Joary eden eLOA-AC Joary eden LOG-AT Joary eden LOW-AT Joary eden LOS-dI IoAry eden IOLPE9OH# // 98Plg []2MxXP/LO-VI1 4L0H-VI PLOH-VI N vod-V I pod-VI q6c-VI P6c-VI IoAry eden IdAry eden IoATy eden IaaAry eden IoAry eden IoAry eden IoArTy eden eIZLIAd -SLI ‘OU DOUdIIJaI YURGUIDH / uoT]edOT sJduIeS ‘SASATVNV dIHV GONV SI] Od SUAAWOAN NOISSAOOY ANVENAD GNV NOILVNYOAN] NOLLOATIOD DUDIJIANDUL * SISUAI]ISDAG * DUDIAIUL{DYIS ° SI]DJUIAPIIIO * IDIPUDJIZ-IDAOU * 1UuOSDUL "JT SISUIUTJOADI “T so1sodsg ‘| alav bs 136 TABLE 2. AFLP PRIMER AND ADAPTER SEQUENCES. Primer AdlEcoRI Ad1Msel prampEcoRI prampMsel FAM-EcoRI HEX-EcoRI MselI + CAA Msel + CAT MselI + CAG novae-zealandiae were excluded from this analysis due to technical difficulties with the DNA extraction from the leaf material. Dilutions of the genomic DNA extract of 1:10 in ultrapure water were used in PCR reactions. The contigu- ous ITSI, 5.88, and ITS2 regions of nuclear ribosomal DNA were PCR-amplified using the primers ITSLEU (Baum et al. 1998) and ITS4 (White et al. 1990) in final reaction volumes of 25 ul. Positive amplifications were purified using the MO BIO UltraClean PCR Clean-up DNA Purification Kit (MO BIO Laboratories, Inc., Solana Beach, CA). Internal primers ITS2 and ITS3 (White et al. 1990) were used in addition to ITSLEU and ITS4 in cycle-sequencing reactions in order to extend fragments and clarify ambigu- ities. Fragments were sequenced with the BigDye 3.1 kit (Applied Biosystems, Foster City, CA) following the manufacturer’s protocols, and visualized using the ABI PRISM 3100 Genetic Analyzer (Applied Biosystems). Sequences were manually aligned using Sequencher 3.1.1 (Gene- Codes Corp., Ann Arbor, MI) and MacClade 4.04 (Maddison and Maddison 2001). AFLP Methods AFLP fingerprinting was conducted follow- ing a modified protocol based on the methods described by Vos et al. (1995). DNA extracts prepared for ITS analysis also were used for this study undiluted. Approximate DNA con- centrations for all samples were estimated to contain a range of concentrations from 10 ng/ul to 50 ng/ul using an ethidium dot test. DNA template of each sample was digested using the infrequent endonuclease cutter EcoR1 and the frequent endonuclease cutter Msel. Immediately following digestion, the entire digestion reaction was combined with an equal volume of ligation mix. The resulting fragmented DNA template con- taining “sticky ends” was diluted five-fold and subsequently amplified by PCR using a pre- selective primer mix. This step effectively reduces the number of possible fragments by approxi- mately 1/16" (Meudt and Clarke 2007). The MADRONO [Vol. 58 Sequence 5’ =CTCGTAGACTGCGTACG— 3” 5 =CGACGATGAGRCCTGAG— 3! 5’ =GACTGCGTACCAATTCA— 3’ 5 “GATGAGTCCTGAGTAAC= 3’ 5’ -GACTGCGTACCAATTCAAC- 3’ 5) =GACT GCG TAGCAATTCACG=3" 5 =GATGAGTCCTGAGTAACAA— 3 5’ -GATGAGTCCTGAGTAACAT= 3’ 5’ -GATGAGTCCITGAGTAACAG— 3” pre-selective reaction condition consisted of 30 cycles of 94° for 30 sec, 56° for 1 min, and 72° for 1 min. Three combinations of selective primer sets were used to produce a final AFLP fingerprint for each sample (Table 2). Each set of selective primers consisted of a primer region matching the known adapter sequence, as well as three selective nucleotides on the 3’ end of the MseI primer and three selective nucleotides plus a florescent label on the 3’ end the of EcoRI primer. Template for the pre-selective PCR was diluted 6-fold and then combined with a master mix containing one set of selective PCR primers. A step-down PCR was used to amplify the selective fragments in a program consisting of 13 cycles of 94° for 30 sec, 65° for 30 sec (—0.7° per cycle), and 72° for 1 min, followed by 24 cycles of 94° for 30 sec, 56° for 30 sec, and 72° for 10 sec. The final selective PCR product fragments containing a fluorescently labeled EcoRI end and unlabeled Msel end were analyzed undiluted using an ABI 3100 genetic analyzer (Applied Biosystems). Initial fragments were sized first using the analysis software GeneScan (Applied Biosystems) and using the program by GeneMar- ker© (SoftGenetics, State College, PA). After a comparison of fragment calling using both programs, all samples were analyzed using GeneMarker®. Fragments were recorded for each sample in a data matrix based on a binary system (1 for presence, 0 for absence); a data matrix was developed for each primer combination and then all data was collated into a single data matrix. To test reproducibility of results, twenty percent of all samples selected at random for each primer pair were re-analyzed, starting with the initial DNA extracts. Fragment peaks that were deter- mined to be consistently low (below 300 peak intensity) or unpredictable were dropped from the matrix table. Data Analysis Phylogenetic analyses of ITS sequences were conducted using Phylip version 3.68 (Felsentein 2004). All characters were weighted equally, 2011] character state transformations were treated as unordered, and gaps were treated as missing data. Most-parsimonious trees were obtained in Phylip using the “‘branch-and-bound” method of exact search implemented by the analysis unit DNA- PENNY. Bootstrap re-sampling (1000 replicates) was used to assess nodal support (Felsenstein 1985). Most parsimonious trees were generated from a search of 100,000 trees and a final tree was derived using a strict consensus tree method (Felsenstein 2004). Additional tree searches were conducted in greater volumes, up to 1,000,000. However, larger searches produced the same final tree, thus a smaller tree search was selected to reduce run-time during bootstrapping. Several combinations of Lilaeopsis species outgroups were explored before selecting L. novae-zelandiae as the outgroup. This selection was based on indications of a potential sister group relationship between L. novae-zelandiae and L. occidentalis, which was supported by ITS phylogenetic anal- ysis of this genus within the Apiaceae tribe described in Downie et al. (2008). Genetic distances to determine branch lengths were calculated in Phylip using the Jukes-Cantor method implemented in DNADIST and a Fitch-Margoliash (FITCH) search. AFLP phylogenetic analysis was performed us- ing the program Phylip version 3.68 (Felsenstein 2004). A genetic distance matrix was created using the techniques described by Nei and Lei (1979) as implemented by RestDist in Phylip (Felsenstein 2004). The output matrix was then input into NEIGHBOR using the UPGMA method of cluster analysis (Felsenstein 2004). Using this approach, an output tree was con- structed by successive clustering using an aver- age-linkage method of clustering. The output file was then plotted as both a rooted and unrooted tree. A search for the most parsimonious trees was implemented first using the branch-and- bound algorithm of DOLPENNY (100,000 trees searched) in Phylip following bootstrap analysis using 100 replicates. A 50% majority-rule con- sensus tree was then generated to condense the results into a final tree, which is presented here. Previous studies of Lilaeopsis using AFLP analysis have not previously been reported. As such, outgroup selection for AFLP parsimony analysis was determined following variable, preliminary analysis replicates. Lilaeopsis schaff- neriana was selected as this species is the closest geographically to both ZL. masonii and _ L. occidentalis. Further, L. schaffneriana also dem- onstrated sufficient genetic differences to be used as an appropriate outgroup. To further visualize potential multi-dimen- sional correlation of AFLP data based on genetic similarities, an additional genetic dis- tance matrix was derived using Manhattan distance (StatisiXL; www.statistixl.com). These FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS [37 data were then analyzed using a _ principal coordinates analysis (PCA) using the Microsoft Excel© add-in program GenAIEx 6.2 (Peakall and Smouse 2006). To determine whether a measurable degree of genetic dissimilarity among L. occidentalis (WA and CA samples) and L. masonii (CA samples) could be attributed to geographic distance, an additional test of molecular variance based on geographic origin as measured by Global Position System (GPS) also was tested. A two-way analysis of variance was assessed for collections of L. occidentalis and L. masonii using an analysis of molecular variance (AMOVA) with GenAIEx 6.2 (Peakall and Smouse 2006). Significance was assessed using 99 permutations. RESULTS DNA sequence analysis of the ITS regions ITS1, 5.88, and ITS2 of nuclear ribosomal DNA, based on a most parsimonious search of 64,631 trees using a branch-and-bound method, revealed no differences between California samples of L. occidentalis and L. masonii samples, including a GenBank accession for L. occidentalis (100 of 100 trees) (Fig. 2). Within the L. occidentalis\L. masonii clade, samples collected from Washing- ton clustered separately. Distance values between Washington and California samples were low for single collections from Lawrence Lake and Mason’s Lake (0.1 and 0.3%, respectively). However, a second sample from Lawrence Lake exhibited higher distance values (1.5%), which may be due to missing data. Distance values for California samples of L. masonii and L. occiden- talis were 0% across all samples. Comparatively, distance values between the additional species used for this study ranged from 1.2—8%. Distance based analysis of ITS sequences found an identical tree structure as the strict consensus tree inferred from most parsimonious results implemented by DNAPENNY. Bootstrap esti- mates from 1000 replicate analyses yielded 100% nodal support for all branches. Branch placement and relationship of Lilaeopsis species used in this study are consistent with results of a previous ITS phylogenetic analyses by Downie et al. (2008), though that study excluded L. masoniti. The three AFLP primer combinations gener- ated 274 unique fragments among 29 samples, with only 21 fragments shared or monomorphic between the five species used in this study. Although a small sample size was used, specifi- cally for L. brasiliensis, L. schaffneriana, and L. mauritiana, the large number of shared fragments is potentially indicative of low genetic diversity within this genus, which is consistent with species exhibiting high morphologic plasticity (Linhart and Grant 1996). The number of total diagnostic bands from the three markers combined data set 138 MADRONO t—{ 0.003 substitutions/base L. brasiliensis2 L. brasiliensis? L. mauritiana L. occidentalis (Bodega Bay, CA) 1A-B04a, Napa L. occidentalis (Mason Lake, WA) L. occidentalis (Lawrence Lake2, WA) L. occidentalis (Lawrence Lake,1 WA L. occidentalis (GenBank) 3 Mile Slough, CA Grizzly Island, CA Brannan Island, CA 1A-H07b, Napa 1A-29a, Napa Collinsville, CA 2W-B02a, Napa 2w-B02b, Napa 2E-A04, Napa McAvoy Harboy, CA Reach 3, Napa 2E-B07a, Napa 2E-A07, Napa 1A-B04b, Napa Liberty Island, CA 1A-H07a, Napa 2W-B02a, Napa 1A-001, Napa 1A-29b, Napa Maxwell Bridge, Napa Twitchell Island, CA 1B-S07, Napa L. carolinensis (GenBank) L.schaffneriana? L.schaffneriana2 L.novae-zelandiae (GenBank) Fic. 2. Strict consensus tree derived from ITS sequence data using the branch and bound method implemented by DOLPENNY. Branch values are Bootstraps. Upper left tree illustrates distance values, branch lengths are proportional to the number of nucleotide substitutions per base. varied between species, ranging from a low of 98 bands for one sample of L. schaffneriana, to a high of 135 bands for the Mason Lake, WA sample of L. occidentalis, with a mean number of AFLP bands equaling 123 (SD = 9.5) (Table 3). Within species genetic distance values (Man- hattan distance calculated from the AFLP matrix) ranged from a low of 1.4 for L. masonii (Napa River site #1A-H07, large and small forms), to a high of 6.6 for L. masonii (Napa site 1A-BO4 and Twitchell Island collections). Between species values ranged from a low of 4.1 for L. occidentalis (Mason Lake, WA) and L. masonti (Napa River #2W-B02), to high of 11.9 for L. schaffneriana and L. occidentalis (Lake Lawrence, WA collection). Principal coordinate analysis (PCA) illustrated an overlapping association between samples of L. occidentalis and L. masonii (Fig. 3), but a clear differentiation between the L. occidentalis/L. masonii cluster and all other Lilaeopsis species examined in this study, i.e., L. schaffneriana, L. mauritiana, and L. brasiliensis. The small separa- tion observed between the L. occidentalis and L. masonii data may be attributed to geographic distance. Results of AMOVA analysis derived 2011] FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS 139 TABLE 3. AFLP FRAGMENT NUMBERS FOR EACH SPECIES. Mean or Total No. of Fragments Species CAA CAG CAT Total L. masonii 33.0 46.1 21.9 125.1 L. occidentalis a paw 50 22:9 128.8 L. schaffneriana subsp. recurva 47.5 34.5 16 99.5 L. brasiliensis a2 39 21 114 L. mauritiana 43 38 16 99 All Fragments 123 from grouping samples based on geographic location indicated that approximately 73% of genetic variation was distributed between groups and thus 23% among groups, supporting the conclusion that most observed genetic variation 1s due to geographic distance (P = 0.01). The UPGMA cluster phenograms provides additional support for the results of the PCA analysis, illustrating a combined grouping of L. occidentalis (CA and WA) and L. masonii samples (Fig. 4). Within the L. occidentalis and L. masonii clade, samples collected from Wash- ington clustered separately from samples col- lected within California, corroborative of results of AMOVA indicating that variation within this clade is due in large part to geographic distance. Samples of L. brasiliensis and L. mauritiana cluster separately but are sister to the L. occidentalis/L. masonii clade; L. schaffneriana samples also cluster separately but are sister to all other specimens/species used for this study. The most parsimonious tree from the maximum parsimony analysis supports a single L. occiden- talis/L. masonii clade; however, additional reso- (gm > L. occidentalis (CA) Lioccidentalis (WA) Coord.2 : | | | : L.masonii lution within this taxon is less certain of the specific placement of Lilaeopsis samples, based on geographic location (Fig. 5). DISCUSSION Taxonomic implications. Within the last two decades, the use of genetic techniques to distinguish discrete evolutionary units has become common place in systematic biology. Use of genetic data in the protection of endangered species when morpho- logical (or other character) information is either unreliable or 1mpossible is just one reason why this approach to species identification and delimitation is SO Important (Avise 2003). Thus, sole reliance on morphological, geographic, reproductive behavior or some combination of non-genetic characters to delimit taxa is no longer defensible when diagnostic genetic information is available and can be readily assessed. In the case of Lilaeopsis masonii and L. occidentalis, neither ITS sequence nor AFLP fragment length data support the recognition of the L. masonii as a distinct evolutionary entity. L.schaffneriana L. mauritiana L. brasilensis Coord.1 FIG. 3. Principal coordinates analysis (PCA) of AFLP fragment data matrix. Codes for Napa collections, e.g., 1A-H07, indicate different collection locations and dates along the Napa River specific to the Napa River Flood Protection Project. 140 MADRONO 39.8 49.5 59 [Vol. 58 9.5 Lawrence Lake earns Lawrence Lake 2 Li/aeopsis 128 occidentalis : Lawrence Lake 3 (Washington) 9.5 Mason Lake 1 Mason Lake 2 1A-29a 1A-B04a Ey 3. 5 A-29b L. masonii (Napa, CA) 1A-H07a 4A-HO7b 3S Maxwell Bridge 2W-B02a 2W-B02b McAvoy Harbor 3 Mile Slough Brannan Island L. masonii Liberty Island (California) Collinsville | Grizzly Island 4{A-001 L. masonii Reach 3 (Napa, CA) 8.3 2E-B07 9.2 __L. occidentalis =e Bodega Head — atifornia) 13. L. masonii 1A-B04b (Napa, CA) 14 : L. masonii Twitchell Island — (California) Lilaeopsis brasiliensis Lilaeopsis mauritiana 2.9 Lilaeopsis schaffneriana1 Lilaeopsis schaffneriana2 Fic. 4. UPGMA Cluster Phenogram (rooted and unrooted trees) from AFLP data matrix of five species of Lilaeopsis. Numbers are distance values. Codes for Napa collections, e.g., 1A-H07, indicate different collection locations and dates along the Napa River specific to the Napa River Flood Protection Project. The morphological and geographic information to support two distinct taxa is weak, ambiguous, and unreliable at best. Fallon’s (2007) arguments regarding the im- portance of using genetic information to resolve taxonomic issues for species protection is borne out in our study. While her review focused solely on vertebrates, and on only those species, infraspecific or population segments proposed for listing, not those already listed, our results add further emphasis for use of molecular techniques in conservation efforts. We concur that multiple genetic markers are essential for a thorough assessment of taxonomic or population unit (or at any appropriate level) when consider- ing of formal protection. We further suggest that use of best available science such as existing or generating new genetic information is equally valid for the periodic reviews of listed species required of both the federal and state agencies. Further and relevant to L. masonii, use of genetic data is likely to be essential during a de-listing review process. Based upon several lines of evidence, including decades of fieldwork throughout the range of L. masonii, observations from the most recent monograph (Affolter 1985), and our molecular genetic analyses, we urge that this rare taxon no longer be recognized as a separate taxonomic entity. Rather, L. masonii should be subsumed within the larger, much more widespread, com- mon, and equally variable species, L. occidentalis. 2011] 96/100 100/100 71/100 96/100 57/100 58/100 FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS 14] Reach 3, Napa A1-001, Napa L. occidentalis (ML1, WA) L. occidentalis (ML2, WA) Maxwell Bridge L. occidentalis (LL3, WA) 1A-29b, Napa 1A-B04a, Napa 1A-29a, Napa 2W-B02a, Napa 3 Mile Slough, CA McAvoy Harbor, CA L. occidentalis (Bodega Hd,CA) L. occidentalis (LL1, WA) 1A-H07a, Napa 1A-H07/b, Napa Brannan Island, CA Grizzly Island, CA 2W-B02b, Napa Collinsville, CA 1A-B04b, Napa L. occidentalis (LL2, WA) 2E-B0/, Napa Twitchell Island, CA Liberty Island, CA L. brasiliensis L. mauritiana L. schaffneriana1 L. schaffneriana2 FIG. 5. Most parsimonious tree from AFLP data matrix of five species of Lilaeopsis. Codes for Napa collections, e.g., LA-H07, indicate different collection locations and dates along the Napa River specific to the Napa River Flood Protection Project. Significant morphological, but limited genetic, variation exists both within and among popula- tions of Lilaeopsis throughout the Pacific Coast of North America, from the Queen Charlotte Islands in British Columbia to the inland islands of the Sacramento-San Joaquin Delta in the Great Valley of California. Importantly, this variation does not follow any consistent environ- mental gradient for either taxon. As such, one intrinsically variable species, not two, of west coast Lilaeopsis should be recognized in relevant floras, including those for North America. Important additional circumstantial support comes from the very wide geographic ranges, some amphitropical, of the great majority of other species of Lilaeopsis, including L. chinensis Kuntze, L. carolinensis, L. shaffneriana, L. macloviana A. W. Hill, and L. novae-zelandiae, among others. A large geographic range is not surprising for all these species, given their vigorous vegetative reproduction by easily frag- mented rhizomes and their restriction to aquatic habitats, many with bi- or multi-directional flow vectors (e.g., Napa River, Sacramento River, Pacific Ocean). Field observations suggest a possible explana- tion for the inter- and intra-populational varia- tion individual ramet size for both western North American Lilaeopsis. Periods of rapid spring growth occur during the spring tides and snowmelt from the Sierra Nevada, when temper- atures warm sufficiently to encourage an increase in photosynthetic activity. This increased vegeta- tive growth occurs when floodwaters from the 142 MADRONO Sacramento, San Joaquin and Napa Rivers are at their height of volume and rate of flow. Thus, the comparatively high kinetic energy of flowing water during the spring run-off, coupled with this species’ preference for river banks and shores characterized by high light and open exposure, combine to restrict vegetative growth to a comparatively shorter plant less vulnerable to being dislodged from its habitat. Relatively taller Lilaeopsis ramets are invariably found compara- tively further from the shoreline in shadier and relatively lower energy microhabitats than com- paratively shorter stature ramets. Aquaria enthu- siasts who work with various species of Lilaeopsis have dubbed the short stature coupled with dense growth phenomenon the “lawn effect” (http:// www. freshwateraquariumplants.com). Conversely, observations of Lilaeopsis species submerged in (low energy) water reveal individual leaves grow comparatively longer. Affolter’s (1985) greenhouse experiments and observations that demonstrated that for least eight of the 13 species Lilaeopsis studied (including L. occidenta- lis), material grown in submerged pots had larger and wider leaves, more septae, and wider rhizome diameter. In his monograph of the thirteen species known in 1985, increased periods of inundation result in a suite of morphological changes, including an increase in leaf length and increases in both peduncle and pedicel lengths (Affolter 1985). Lastly, the rejection of leaf length as a key diagnostic character distinguishing two otherwise very similar taxa has precedent in Affolter’s lumping of all Andean, Fuegian, and Patagonian material into a single species, L. macloviana, synonomizing thirteen previously described taxa. Regulatory implications. Neither CESA nor the federal ESA, as amended, protects any vascular plant distinct population segment as does the ESA for specific vertebrate populations. While an argument can be made that this is a form of taxon chauvinism, plant species are not protected below the infraspecific level. Such a comparison is important, because some vertebrate species that were listed relatively soon after the ESA was passed have since been determined not genetically distinct from common widespread relatives, but they continue to be formally protected because of the DPS provisions. For example, the San Francisco garter snake (Thamnophis sirtalis tetra- taenia), a highly restricted taxon in central coastal California, was determined, through an exami- nation of the clade’s mtDNA (Janzen et al. 2002), to be a member of a California clade of the widespread common garter snake. These authors concluded that morphologically based subspecies designations of 7. sirtalis in western North America were invalid because they did not reflect reciprocal monophyly of mtDNA _ sequences. [Vol. 58 Extrapolating Janzen et al.’s (2002) logic to our genetic work with Lilaeopsis, the parallel conclu- sion that the specific designation of L. masonii is invalid is compelling. Because neither the CESA nor the ESA include DPS provisions for plant species, L. masonii no longer warrants protection as a State “rare” species and the allocation of limited recovery resources. Given the widespread nature of Lilaeopsis occidentalis + L. masonii, and the large number of projects (both existing and proposed) requiring mitigation and monitoring of the rare L. masonii, a timely review of our findings is essential. Conservation dollars are few, and they should be applied to truly rare, threatened, and real discrete species. Finally, Pavlik (2003) recently examined the role of state- and federally-listed species protect- ing the ecosystems in which they are found. Of relevance is the notion that some _ protected species provide a “regulatory umbrella” for other species that are unlisted, but are rare, in decline, or otherwise of conservation concern. Lilaeopsis masonii has long served to restrict, prevent, or slow the conversion, degradation or destruction of wetlands throughout the Sacramento-San Joaquin Delta, Suisun Marsh, and San Francisco Bay ecosystems, thereby protecting associated but unlisted species of conservation concern. While a suite of other protected wetland plant taxa exist in these ecosystems (e.g., Cirsium hydrophilum Jeps. var. hydrophilum [Suisun this- tle], Cordylanthus mollis A. Gray ssp. mollis [soft bird’s beak], etc.), there are many more that are rare, in decline, and not listed (e.g., Cicuta maculata Lam. var. bolanderi (S. Watson) G. A. Mulligan [Bolander’s water hemlock], Plantago elongate Pursh [slender plantain], Lycopus asper Greene) (see Baye et al. 2000). Thus we acknowl- edge that a delisting of Mason’s lilaeopsis may further expedite wetland habitat loss in central California. Nonetheless, conservation in the twenty-first century demands the use of best available science, despite the unintended conse- quences that may occur. Ultimately, government agencies charged with the protection of our biodiversity must redouble their efforts to em- brace new scientific results that affect listed species, commit to diligent review of listed and candidate species, and disseminate accurate and up-to-date information. Similarly, conservation- ists should redouble their efforts to provide the best available science for decision-making. The time to embrace current molecular genetic techniques in routine conservation decision-mak- ing has come. ACKNOWLEDGMENTS The authors would like to thank the Napa County Flood Control and Water Conservation District, particularly R. Thomasser, for funding this work; L. 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WSP ENVIRONMENTAL STRATEGIES (WSP). 2007, Year 7 (2007) Rare plants of the lower Napa River/Napa Creek Flood Protection Project. Final Monitoring Report prepared for the Napa County Flood Control and Water Conservation District, Napa, CA. ZEBELL, R. K. AND P. L. FIEDLER. 1996. Final Report. Restoration and recovery of Mason’s lilaeopsis. Phase II. Report to the Shell Oil Litigation Settlement Trustee Committee and the Endangered Plant Program, Natural Heritage Division, Cali- fornia Department of Fish and Game, Sacramento, CA.Website http://nrm.dfg.ca.gov/FileHandler. ashx?DocumentVersionID=3709 [accessed 28 July 2011). MADRONO, Vol. 58, No. 3, pp. 145—152, 2011 A COMPARISON OF THE EFFECTS OF NA>;SO, AND NACL ON THE GROWTH OF HELIANTHUS PARADOXUS AND HELIANTHUS ANNUUS (ASTERACEAE) M. O. MENDEZ! AND O. W. VAN AUKEN Department of Biology, University of Texas at San Antonio, San Antonio, TX 78249 monica.mendez@tamiu.edu ABSTRACT Helianthus paradoxus Heiser (Asteraceae, puzzle sunflower), is a federally threatened hybrid species found in salt marshes of west Texas and New Mexico. Helianthus annuus L. (Asteraceae, common sunflower) is one of the parent species and is found throughout North America, but it is not present in the inland salt marshes where H. paradoxus is found. Helianthus paradoxus has previously been described as a halophyte, but its tolerance to Na »SOx4, one of the major salts found in its habitat, has not been investigated. However, salinity has been identified as a major abiotic factor influencing the limited distribution of H. paradoxus populations. In this greenhouse study, the effects of elevated concentrations of NasSO,4 and NaCl, at equal ionic strengths (0.00, 0.09, 0.17, 0.34, and 0.51), on the survival and dry mass of both H. paradoxus and H. annuus were examined. In the three-way factorial experiment, the effects on dry mass observed were dependent on the species, the type of salt and the ionic strength of the salt. Helianthus paradoxus produced more dry mass than AH. annuus in both salt treatments; however, NaCl was more inhibitory of dry mass production for both species with plants unable to survive the highest salt treatments. While dry mass of H. annuus decreased with increasing ionic strengths of both salts, dry mass of H. paradoxus increased by 38 to 72% in low to moderate ionic strengths of NazSO, relative to the nonsaline treatment. Both species were less tolerant of NaCl than Na,SO, with A. paradoxus seeming to have moderate and high tolerance to elevated Cl” and SO,’ ionic strength, respectively, while H. annuus had low to moderate tolerance. Greater dry mass production in Na»,SOxq, along with tolerance to both salts, suggests that low to moderate sulfate soil salinity will enhance the dry mass production of H. paradoxus. Key Words: Halophyte, Helianthus annuus, helianthus paradoxus, ionic strength, NaCl, Na>»SOxg, salt tolerance, sunflower. Helianthus paradoxus Heiser (Asteraceae, puz- zle sunflower) is a federally threatened species with limited distribution in salt marshes in west Texas and New Mexico (Correll and Johnston 1979; Poole and Diamond 1993; McDonald 1999). Hybridization studies (Heiser 1958, 1965; Abbott 1992) and molecular analysis (Rieseberg et al. 1990; Rieseberg 1991; Rieseberg et al. 1991) have determined that H. paradoxus is a stabilized hybrid species between H. annuus L. (Asteraceae, common sunflower) and H. petiolaris Nutt. (plains sunflower). Although H. paradoxus shares several morphological and ecophysiological traits with its parental species (Rosenthal et al. 2002), it has diverged and is genetically isolated from its progenitors and considered a separate species. Helianthus anuus is common throughout North America and grows in disturbed, heavy clay soils that are moist in the spring and dry out by mid- summer. Helianthus petiolaris is found in dry, sandy soils in western North America, while H. paradoxus grows in heavy, waterlogged, saline soils (Van Auken and Bush 1998). ‘Current address: Department of Biology & Chem- istry, Texas A&M International University, 5201 University Boulevard, Laredo, TX 78041, USA. Ecological and ecophysiological studies of H. paradoxus have determined that this homoploid hybrid species is salt tolerant, unlike its parental species. Helianthus paradoxus 1s restricted to inland salt marshes with salt levels of approxi- mately 10 g kg ' (Poole and Diamond 1993; U.S. Fish and Wildlife Service 2005; Grunstra and Van Auken 2007a, b), while H. annuus and H. petiolaris can be found in low saline soils (<0.02 g kg™' soil sodium, Welch and Rieseberg 2002). Helianthus paradoxus is a better compet- itor than its progenitors in saline soils (Bush and Van Auken 2004). The west Texas and New Mexico salt marshes were key habitats in isolating hybrids (Abbott 1992). The parental species are glycophytes and cannot survive the same habitat as H. paradoxus, where other salt tolerant plants are generally present (Poole and Diamond 1993: Lexer et al. 2003: U.S. Fish and Wildlife Service 2005). Additionally, H. para- doxus is capable of sequestering higher sodium and sulfur concentrations and produces greater leaf succulence compared to its parental species while maintaining significantly greater fitness when grown in elevated NaCl concentrations (Welch and Rieseberg 2002) or in field-like highly saline soil conditions (Karrenberg et al. 2006). In the largest known population of H. para- doxus, at the Diamond-Y Spring Preserve near 146 Fort Stockton, Texas, the distribution of JH. paradoxus is mainly affected by soil salinity and soil moisture gradients when biotic factors are not considered (Bush and Van Auken 1997; Van Auken and Bush 1998; Bush 2006a, b). This large desert spring and associated salt marsh has had 1.44 to 2.70 million H. paradoxus plants (Van Auken and Bush 1998), depending on environ- mental conditions. In addition, there are several rare and federally endangered invertebrates found in the marsh (McDonald 1999). Helianthus paradoxus plants consistently establish parallel to the drainage of Leon Creek but their proximity depends on seasonal climatic conditions influencing soil salinity levels and soil water content, both of which decrease dramatically upland from the lowest point in the salt marsh (Van Auken and Bush 1998; Grunstra and Van Auken 2007a, b). When annual rainfall is high, H. paradoxus can be found further from the drainage compared to drier years when plants are located closer to the drainage. The population of H. paradoxus at the Diamond-Y Spring Preserve seems to be dependent on low to intermediate salinity levels and intermediate moisture levels; however, salinity appears to be the major abiotic factor affecting the local distribution of H. paradoxus (Bush 2006a, b). Previous studies have identified H. paradoxus as a salt tolerant species with characteristics of halophytes (Welch and Rieseberg 2002; Bush and Van Auken 2004). However, these studies were done with NaCl, one of the major salt compo- nents of the marsh, but not Na»,SO, which is also found at high concentrations in soils of its salt marsh habitat (Boghici 1997). In addition, the effects of a limited range of salinity levels have been examined. Chloride is generally more toxic than SO,°- and even Na‘ at lower concentrations (Manchanda et al. 1982; Marschner 1995; Frank- lin and Zwiazek 2004; Munns and Tester 2008); however, SO, salinity can be more growth inhibitive than Cl” for some halophytes (Warne et al. 1990; Egan and Ungar 1998). At the same time, separating specific ion effects is difficult and differential effects of Cl” and SO,’ salinity on H. paradoxus are still unknown; consequently, the influence of these anions on the ecological isolation and distribution of H. paradoxus has not yet been elucidated. Ionic strength rather than salt concentration was used as a treatment variable in the present study. It is used as a normalization procedure and was required because of different numbers of ions present in equal molar solutions of NaCl (2 ions) and Na>SO, (3 ions). Therefore, it was important to compare the two salts at concentrations that allowed osmotic potential of the corresponding treatments to be equal. Concentrations of the two salt treatments were based on equal ionic strengths calculated using procedures in Barrow (1966). MADRONO [Vol. 58 TABLE 1. IONIC STRENGTHS OF EACH TREATMENT AND CORRESPONDING SALT CONCENTRATIONS IN G:-KG ' EXAMINED IN THE STUDY. NaCl Na»SOx,4 Ionic strength (g kg~') 0.00 0 0 0.09 eS) 4.1 0.17 10 8.1 0.34 20 16.2 ee | 30 24.3 The objective of this study was to examine the survival, growth and the salt tolerance of ZH. paradoxus in elevated levels of both Na»SO, and NaCl, at equal ionic strengths. Both of these salts are major contributors of the soil salts of the Diamond-Y Spring Preserve. Our experiment also included the more salt tolerant of its parental species, H. annuus, a known glycophyte. MATERIALS AND METHODS Helianthus paradoxus seeds were collected from the Nature Conservancy’s Diamond-Y Spring Preserve near Fort Stockton, Texas (31°00.54’N, 102°55.49’W) and stored dry at 25°C until used. Helianthus annuus seeds were purchased from Native American Seed Farm (Junction, Texas 76849) and stored dry at 4°C. Seeds of both species were cold stratified in Ziploc® storage bags lined with paper towels, wet with deionized water, and maintained at 4°C in the dark for 4 weeks (H. paradoxus) or 8 weeks (H. annuus). To prevent osmotic shock, groups of seeds (approximately 400 seeds for each treatment) were germinated on paper toweling saturated with deionized water (0.00 osmotic strength) in plastic storage bags at 25°C. Germinated seed- lings were transferred to equal or increasing levels of the appropriate salt (NaCl and Na ,SOy, at ionic strengths of 0.00, 0.05, 0.09, 0.13, 0.17, 0.26, 0.34, 0.43, and 0.51) every 2 d until placed in the final salt (NaCl or Na>»SO,) and ionic strength to be tested (0.00, 0.09, 0.17, 0.34, and 0.51). Seedlings were kept in the appropriate solution for a total of 18 days prior to transplanting to pots in the greenhouse containing the corre- sponding treatment (Table 1). For each species, five plants per pot with five replicate pots per treatment (25 plants per species per treatment) were grown for 62 d. Plastic pots (15 cm diameter < 15 cm height) lined with a Ziploc® storage bag (to retain water, salts, and nutrients) were filled with 1.4 kg of air-dried, sieved (5.8 mm mesh screen) soil. The soil was the upper 10 cm of a low nutrient Patrick-series Mollisol (clayey-over-sandy, carbonatic-thermic, typic calciustoll), collected from northern Bexar Co., Texas (Taylor et al. 1966; Van Auken and Bush 1998). The soil was friable, allowing root 2011] TABLE 2. MENDEZ AND VAN AUKEN: SALT TOLERANCE OF TWO HELIANTHUS SPECIES 147 THREE-WAY ANOVA RESULTS FOR THE TOTAL DRY MASS (G) OF TWO SPECIES (HELIANTHUS PARADOXUS AND H. ANNUUS) GROWN IN TWO SALTS (NACL AND NA>SQOq) AND AT FIVE IONIC STRENGTHS (0.00, 0.09, 0.17, 0.34, AND 0.51). All main effects and interactions were significant. “ Significant interactions are designated (* = P < 0.05; ** = P < 0.01; *** = P < 0.001; **** = P < 0.0001). OQ. mh Source* Species Salt Strength Species < salt*** Species < strength** Salt X strength**** Species X salt < strength* Error Total Oo HH Hee Nome.<) extraction and recovery (Bush and Van Auken 2004). Appropriate amounts of anhydrous salts (Na>SO,q, or NaCl) and a single nutrient applica- tion (0.05 g of P from Na,HPO, - 7H,O, 0.07 g N from NH4NOs3, 0.07 g of K from KCl, and 0.03 g of S from MgSO, 7H,O; Tiedemann and Klemmedson 1986) were added to each pot and thoroughly mixed. Before seedlings were planted, 400 ml of deionized water was added to each pot. Thereafter, soil moisture was maintained at approximately field capacity with distilled water. This study was conducted in a fiberglass green- house in which the daytime temperatures ranged from approximately 26° to 38°C and light levels were approximately 36% of full sunlight with a mean photosynthetically active photon flux den- sities of 562 + 135 umol m * s_' measured with a Li-Cor® LI-188 integrating quantum sensor. Plant survivorship was assessed at 62 d for each treatment combination (species X< salt xX ionic strength). Percent survivorship is based on the mean number of erect and green plants out of five plants per replicate pot per treatment. Shoot and root dry mass were measured at the end of the experiment. For shoot dry mass, plants from each pot were clipped at the soil surface and placed in a pre-weighed paper bag. For the ash- free root dry mass, all of the soil and particulate matter from each pot was carefully washed off the roots. Roots were then wrapped in pre- weighed aluminum foil. All plant material was dried in a forced air oven at 90°C to a constant mass. After drying, roots were ashed in a programmable muffle furnace (Fisher Scientific Isotemp®, Fisher Scientific Research, Pittsburgh, PA) at 625°C for 3 hours to obtain the ash-free root dry mass (Bohm 1979). Total dry mass was also calculated by combining the shoot dry mass and ash-free root dry mass per pot. All dry mass data was analyzed using SAS Statistical software (SAS Institute 1999) with each pot as the unit of replication and P = 0.05 as the criterion for significance. Any plant mortalities because of a treatment effect were taken into account by including a dry mass of zero. To SS F Pp 4.11 35.69 <0.0001 14.75 128.17 <0.0001 5.00 43.44 <0.0001 Lo? 12.05 0.0008 0.35 4.78 0.0016 1.67 14.50 <0.0001 0.40 3-5) 0.0108 921 57.05 determine the effects of salt type and ionic strength on the dry mass of each species, a three-way ANOVA including interactions was employed. When significance was detected with the overall ANOVA, two-way and one-way ANOVAs were used followed by Duncan’s Multiple Range Test to examine significant differences between all possible combinations of salt type and salt concentration for each species separately. Mean mortality was calculated, but was not analyzed statistically. RESULTS All H. paradoxus and H. annuus plants died in the highest ionic strengths of NaCl examined (0.34 and 0.51). However, both species demon- strated 100% survivorship in the no-salt treat- ment. For all Na»,SO, treatments, 100% of H. paradoxus plants survived, while 100% of the H. annuus plants survived in each of the Na»,SO, treatments except at the 0.51 ionic strength, the highest Na»SO, concentration tested. Plant sur- vivorship for H. paradoxus in NaCl treatments was 100% at the 0.09 and 0.17 ionic strengths, while 80% and 40% of H. annuus plants survived in these same ionic strengths, respectively. Three three-way ANOVAs were used to analyze shoot, root, and total dry mass; however, only total dry mass data will be presented. Results were similar for mean shoot and root dry mass (analysis not shown). The three-way ANOVA (Table 2) demonstrated a significant overall species, salt, and ionic strength effect on total dry mass. In addition, the three two-way interactions were significant as was the three-way interaction. To demonstrate more clearly the experimental results, two of the two-way interac- tions will be presented first. Overall, H. para- doxus produced more dry mass than H. annuus in both salts (Fig. la, species < salt interaction). For both species, more dry mass was produced in the Na>»SO, treatment than in the NaCl treat- ment. In addition, ionic strength was significant with more dry mass in the lower treatments; & (oe) ] a) GH. paradoxus [_] H. annuus Total (Shoot + Root) dry mass (g pot’) Salt FIG. 1. MADRONO [Vol. 58 H. paradoxus H. annuus Total (Shoot + Root) dry mass (g pot*') 0.17 0.34 0.51 lonic Strength Two-way interaction plots of total dry mass (bars) as the response variable for (a) salt (NaCl and Na,SOx,) by species (Helianthus paradoxus and Helianthus annuus) and (b) ionic strength (0.00—0.51) by species. A three-way ANOVA determined there were significant differences (P < 0.0001) between salt treatments and ionic strengths for each species. Different letters indicate significant differences between means (Duncan’s Multiple Range Test). Lines above the bars represent + one SD (standard deviation). however, dry mass of H. annuus was significantly lower at an ionic strength of 0.09 (Fig. 1b, species xX 1onic strength interaction). The salt xX ionic strength interaction has not been presented because the results can be seen within the three way interaction figure. The significant three-way interaction indicated that dry mass was depen- dent on species, salt type, and ionic strength (Table 2, Fig. 2a, b). Helianthus paradoxus dry mass was higher in Na,SO4 compared to NaCl (Fig. 2a), with the greatest dry mass at the mid ionic strengths (Fig. 2a). On the other hand, H. annuus dry mass was lower than AH. paradoxus, but H. annuus did produce more dry mass in NasSO, compared to NaCl (Fig. 2b). In addi- tion, as the ionic strength increased dry mass of H. annuus decreased. In comparison to the Na»,SO, treatments, dry mass of both species was significantly reduced in NaCl at elevated ionic strengths (Duncan’s Multiple Range Test, P < 0.0001, Fig. 2a, b). In the no-salt treatment, growth of H. annuus was elevated compared to AH. paradoxus (not signifi- cantly). However, for all levels of salt addition, total dry mass of H. paradoxus was greater than H. annuus, yet the differences were salt depen- dent. For both species, mean total dry mass was elevated in Na>,SOy, at an ionic strength of 0.09 compared to NaCl. Dry mass of H. paradoxus was 32% greater than dry mass of H. annuus in this treatment. In the 0.17 to 0.51 ionic strengths of Na>SOxq, mean total dry mass of H. paradoxus was 2- to 12-fold greater than H. annuus. Both species produced less dry mass in NaCl treat- ments; however, H. annuus dry mass was reduced most by NaCl. Total dry mass of H. paradoxus was 7-fold greater than H. annuus in the 0.09 NaCl treatment. This difference between species in NaCl increased at the 0.17 and 0.34 ionic strengths of NaCl where dry mass of dH. paradoxus was 15-fold greater than H. annuus. DISCUSSION Salt tolerance is the ability of a species to grow and adjust to the presence of a specific ion (ionic effect) or to adapt to the general effects of low water potentials (osmotic effect) (Ungar 1991). In this study, a potential ionic effect and an osmotic effect were investigated using NaCl and Na,SO,4 salts at increasing ionic strengths. Both salts are found in HA. paradoxus habitats at various concentrations (Boghici 1997; Van Auken and Bush 1998; Lexer et al. 2003). To differentiate between the effects of the two salts and the Cl™ and SO,” anions, equal ionic strengths were used in the separate salt treatments (Barrow 1966). Due to greater salt tolerance of H. annuus in NaCl and mixed salt environments compared to H. petiolaris (both purported parental species) (Ashraf and Tufail 1995; Welch and Rieseberg 2002; Bush and Van Auken 2004; Karrenberg et al. 2006; DiCaterina et al. 2007), H. annuus was used as a comparative species for salt effects on growth of H. paradoxus. As in previous studies, this investigation demonstrated that H. paradoxus was more salt tolerant than one of its parental species, H. annuus, and even produced slightly more dry mass in low saline soils compared to H. annuus (Figs. 1b, 2a, and 2b). Helianthus paradoxus produced 70% more dry mass than H. annuus over all treatments (data not shown) and consistently produced more dry mass in elevated salinity soils (Fig. 2a, b), especially in the sulfate treatments. Results of this greenhouse study are consistent with observations that H. paradoxus has greater fitness in saline conditions than H. 2011] nad ° a) H. paradoxus N n N ° Total (Shoot + Root) dry mass (gP°) Oo on lonic Strength mmm Na2so4 C— NaCl b) H. annuus No oi So ~ ra) = So Total (Shoot + Root) dry mass (gP°) oO —_ an an = o lonic strength Fic. 2. Three-way interaction plots of total dry mass (bars) as the response variable for (a) Helianthus paradoxus and (b) Helianthus annuus. Salts were Na 2SO, (black bar) and NaCl (gray bar) at ionic strengths of 0.00—0.51. A three-way ANOVA deter- mined there were significant differences (P < 0.0001) between salt treatments and ionic strengths for each species. Different letters indicate significant differences between means (Duncan’s Multiple Range Test) within a species. Lines above the bars represent + one SD (standard deviation). annuus (Welch and Rieseberg 2002; Bush and Van Auken 2004; Karrenberg et al. 2006), and that H. annuus is not expected to be found in areas with elevated soil salinity where H. para- doxus is able to grow and outcompete H. annuus and probably other non-halophytic species (Ab- bot 2003; Van Auken and Bush 2006). Both Helianthus spp. exhibited a specific ionic growth inhibition at elevated levels of Cl, compared to SQ,’ , and differences in salt tolerance between species were evident. For both species, NaCl caused plant mortality at ionic strengths of 0.34 and 0.51, yet H. paradoxus was MENDEZ AND VAN AUKEN: SALT TOLERANCE OF TWO HELIANTHUS SPECIES 149 more tolerant than H. annuus to low to moderate NaCl concentrations (Fig. 2). Greater dry mass production and survivorship of H. paradoxus plants compared to H. annuus in NaCl treatments was also observed by Welch and Rieseberg (2002) in corresponding NaCl treatments (100 and 200 mmol LL! [~6 and 12 g kg '] NaCl). Plant survivorship of H. annuus indicated that low Na»SO, levels seem to be less inhibitive than NaCl but significant dry mass reduction was still observed above 0.09 ionic strength treatments. Dry mass of H. paradoxus, on the other hand, was enhanced by low to moderate levels of Na>»SO,4 with 38 to 72% greater dry mass, relative to the no-salt treatment. This corresponds to field observations where H. paradoxus was most abundant in soil salinities (mainly Na, K, Ca, and Mg chlorides with less SO4) ranging from 5 to 12 g kg '' in the Diamond-Y Spring Preserve (Boghici 1997; Van Auken and Bush 1998; Bush 2006b; Grunstra and Van Auken 2007a, b). Although previous studies did not differentiate between the effects of NaCl and Na>,SO, (Welch and Rieseberg 2002; Bush and Van Auken 2004; Karrenberg et al. 2006; Van Auken and Bush 2006), those studies support the salt tolerance of H. paradoxus to low concentrations of NaCl. Results consistently demonstrated that biomass of H. annuus was statistically reduced by NaCl alone (Welch and Rieseberg 2002) or by low levels of mixed salts (Na‘*, Cl, and SO,” included; Bush and Van Auken 2004; Karrenberg et al. 2006), while H. paradoxus demonstrated lower productivity in NaCl alone and greater growth and productivity in the presence of SO4*- as observed in the present study. Greater salt tolerance of H. paradoxus to NaCl, as compared to H. annuus, has been attributed to significantly greater fitness along with Na* accumulation, leaf succulence, and water use efficiency (Welch and Rieseberg 2002). Because the salts were at equal ionic strengths and Na* seems to serve as an osmoticum for H. paradoxus (Welch and Riese- berg 2002), it can be assumed that Cl is causing reduced productivity in comparison to SO,’ . Molar concentrations of Cl” in the soil water of the Diamond-Y Spring Preserve are approxi- mately 1.5 times that of molar concentrations of SO, ; therefore, Cl- has the potential of inhib- iting growth of H. paradoxus in its salt marsh habitat (Boghici 1997). However, these data in conjunction with previous salt studies (Welch and Rieseberg 2002; Bush and Van Auken 2004; Karrenberg et al. 2006) indicate that the presence of soil sulfate may have played an important role in the selection for and adaptation of H. paradoxus to the Na-Cl -SO,°~ rich environ- ments. Further, poor tolerance to chloride and sulfate by H. annuus has limited its establishment and therefore, fitness in the H. paradoxus salt marsh habitat. 150 It should be noted that in order to maintain equal osmotic potentials between treatments, molar concentrations of Na* in NaCl treatments were 3.0 times that of Na»,SO, treatments. Nevertheless, the molar equivalents of Na™ at 0.09 and 0.17 ionic strengths were between that of the 0.34 and 0.51 ionic strengths of NasSO4 where dry mass production of H. paradoxus was not inhibited. Since an inert osmotic medium such as polyethylene glycol was not examined, an osmotic effect in combination with an ionic effect cannot be excluded (Katembe et al. 1998; Munns and Tester 2008). In the limited studies comparing phytotoxicity of both salts, greater toxicity to NaCl compared to Na,SO,4 has been demonstrated for other salt tolerant glycophytes and halophytes (Manchanda et al. 1982: Curtin et al. 1993; Franklin and Zwiazek 2004; Pagter et al. 2009). Chloride is more toxic to plants than sulfate possibly due to synergistic phytotoxicity effects with Na‘, differ- ential inhibition of enzyme activity, reduction in plant productivity, and imbalance of nutritional status (Greenway and Munns 1980; Manchanda et al. 1982; Curtin et al. 1993; Wang et al. 1997; Veira Dos Santos and Caldeira 1999; Franklin and Zwiazek 2004). Ion toxicity is dependent on whether the plant possesses adaptations to tolerate the osmotic stress and to exclude and/ or compartmentalize the ion. Although not yet documented, it is possible that H. paradoxus accumulated Cl” along with Na* and may even be more sensitive to Cl’) compared to SO,?” due to poor compartmentalization into vacuoles (Green- way and Munns 1980; Flowers et al. 1986; Munns 1993; Rajakaruna et al. 2003). Chloride may be considered more toxic sometimes because of poor salt tolerance response and thus, high accumula- tion of Cl” over Na’, or in this case SO,” . Toxic cytoplasmic Cl” concentrations have not yet been determined but are assumed to be equal to or slightly lower than Nat (Flowers et al. 1986; Greenway and Munns 1980; Munns and Tester 2008). Sulfate may be required for salt tolerance in H. paradoxus. Tissue ion concentrations were not examined in this study, but previous work suggested that (as in other halophytes) SO,’ , along with Na‘, may be an important vacuolar osmoticum in plant tissue (Greenway and Munns 1980; Karrenberg et al. 2006; Johnston 2006). Leaf sodium and sulfur concentrations were shown to be inversely related to calcium, magnesium, and potassium concentrations but positively related to biomass and succulence in H. paradoxus (Karrenberg et al. 2006). Interestingly, several studies have found a correlation between plants inhabiting waterlogged sulfate rich soils and the presence of vacuole stored flavonoid sulfates which may serve to detoxify excess sulfates alone or in combination with sodium MADRONO [Vol. 58 (Harborne 1975; Tomas-Barberan et al. 1987; Rajakaruna et al. 2003). A possible role of sulphur in the salt tolerance of halophytes also includes the production of methylated sulfonium compounds that accumulate in the cytosol as osmotically compatible organic solutes for com- partmentalization of Na* and Cl” in vacuoles. These organic solutes also serve to detoxify sulfides in salt marsh plants (Flowers and Colmer 2008). Flavonoid sulfates or methylated sulfoni- um compounds have not been measured in tissues of H. paradoxus; however, NasSO, tolerance of this species together with Na* and SO,?~ accu- mulation suggests a possible detoxification mech- anism not yet identified. Ecological isolation of H. paradoxus within the inland salt marshes of west Texas and New Mexico may in part depend on the elevated levels of soil salinity found in soils of these habitats (Boghici 1997; McDonald 1999; Van Auken and Bush 1998, 2006; Abbott 2003). Halophytes are limited to saline environments because of an advantageous adaptation to excess salts and a reduction of competitive ability in non-saline environments (Ungar 1991). Distribution of ZH. paradoxus plants appears to be dependent on low to moderate soil salinity levels at the Diamond-Y Spring Preserve where Na*, Cl” and SO,’ are the prevalent salts from groundwater discharge, while Ca**, Mg**, and HCO3° are secondary ions (Boghici 1997; Van Auken and Bush 1998; Bush 2006b; Grunstra and Van Auken 2007a, b). In the present study, H. paradoxus outperformed H. annuus in both soil salt treatments, partially explaining the inability of H. annuus to survive the salt marsh habitat of H. paradoxus (Lexer et al. 2003; Bush and Van Auken 2004). The presence of salts excluded H. annuus from colonizing the salt marsh (Abbott 2003; Lexer et al. 2003), while the salt tolerance of JH. paradoxus to both NaCl and NazSOy, has allowed this species to establish in the Diamond-Y Spring Preserve and other salt marsh environments in west Texas and New Mexico. Further, tolerance to high concentrations of NasSO4, as demon- strated by significantly greater productivity rela- tive to non-saline conditions, suggests that H. paradoxus may experience a physiological stress response without elevated Na»SO,, which is necessary for optimum growth and _ perhaps necessary for salt tolerance (Munns and Tester 2008). The establishment of H. paradoxus in the Diamond-Y Spring Preserve has been promoted by the selection for transgressive phenotypes promoting salt tolerance (sodium exclusion, calcium uptake, and leaf succulence), as demon- strated by H. paradoxus, along with the presence of specific ions (Na*, Ca**, and SO,’~) in the salt marsh habitat (Abbott 2003; Lexer et al. 2003; Karrenberg et al. 2006). In addition, both parental species are poor competitors in field-like 2011] elevated soil salinity conditions, while H. para- doxus iS a poor competitor against H. annuus in nonsaline conditions (Bush and Van Auken 2004). The potential for hybridization is still present and environmental conditions isolating hybrid Helianthus spp. from parental genotypes appears to have been an important factor in their adaptive evolution of greater fitness in their respective habitats (Lexer et al. 2003; Whitney et al. 2010; Donovan et al. 2010). Populations of H. annuus are found in disturbed pockets of isolated deep, nonsaline soil. Helianthus petiolaris, the other reported parent species to H. paradoxus is also found in some isolated, low saline, sandy soils. However, environmental factors such as the soil conditions required for the isolation and survival of some of the H. annuus X H. petiolaris hybrids was and continues to be the saline soils sur- rounding the isolated springs in this area of west Texas and New Mexico (Abbott 2003; Lexer et al. 2003). The unique salt tolerance of H. paradoxus compared to its parental species along with establishment in the Na‘t-Cl -SO,° domi- nated salt marshes will continue to promote the ecological isolation of H. paradoxus. ACKNOWLEDGMENTS Some support for this project was provided by the Nature Conservancy of Texas, the University of Texas at San Antonio, the Texas Department of Parks and Wildlife, the U.S. Fish and Wildlife Service, and Dr. Loyce and Mr. William Collenback. We especially thank Mr. John Karges of the Nature Conservancy of Texas for help and advice during various stages of this project. LITERATURE CITED ABBOTT, R. J. 1992. Plant invasions, interspecific hybridization and the evolution of new plant taxa. Trends in Ecology and Evolution 7:401—405. . 2003. Sex, sunflowers, and speciation. Science 301:1189-1190. ASHRAF, M. AND M. TUFAIL. 1995. Variation in salinity tolerance in sunflower Helianthus annuus L. Journal of Agronomy and Crop Science 174:351-362. BARROW, G. M. 1966. 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RANDELL, AND L. H. RIESEBERG. 2010. Adaptive introgression of abiotic tolerance traits in the sunflower Helianthus annuus. New Phytologist 187:230—239. MADRONO, Vol. 58, No. 3, pp. 153-184, 2011 THE DIVERSITY AND BIOGEOGRAPHY OF THE ALPINE FLORA OF THE SIERRA NEVADA, CALIFORNIA PHILIP W. RUNDEL Department of Ecology and Evolutionary Biology and the Institute of the Environment and Sustainability, University of California, Los Angeles, CA 90095 rundel@biology.ucla.edu ABSTRACT The alpine zone of the Sierra Nevada of California, defined as non-forested areas at or above 3500 m, includes 385 species (409 taxa) of native vascular plants. Were the alpine boundary defined as at or above 3300 m, the alpine flora would grow to 536 species (570 taxa). There are 97 species that reach elevations of 4000 m and 27 species that reach to 4200 m. Over half of the alpine species occur in just six families, led by the Asteraceae (55 species, 59 taxa), Poaceae (39 species, 47 taxa), Brassicaceae (34 species), and Cyperaceae (31 species). The largest genus present is Carex with 29 species, and 18 more species would be added by lowering the alpine boundary to 3300 m. Next in size are Draba (14 species) and Lupinus (11 species, 16 taxa). Life forms of the flora are heavily dominated by broad- leaved erect perennials (50%), followed in importance by graminoid perennials (21%) and mats and cushions (11%). Annuals and woody shrubs each account for about 6% of the flora. Only nine species are obligate alpine taxa with a range restricted to elevations of 3500 m or above. An additional 67 species (17% of the flora) occur in both subalpine and alpine habitats but not lower. More than a quarter of the alpine species have elevational ranges that extend as low as foothill habitats defined as occurring below 1200 m. In terms of biogeographic affinities, the broad relationships of the flora include the cordillera of western North America (35%), Intermountain region of the Great Basin (20%), Sierra/Cascade axis (16%), and widespread distributions (14%). There are 36 species in the alpine flora endemic to the Sierra Nevada, and another 31 species that are Californian endemics. Key Words: Alpine, arctic-alpine flora, cushion plant, Sierra Nevada, treeline. How large and diverse is the alpine flora of the Sierra Nevada in California and what are its biogeographic relationships? There has been a long history of floristic and ecological studies of the alpine region of the Sierra Nevada addressing this and related issues, but a clear answer to the question has not been achieved. Unlike the majority of alpine regions in the northern hemisphere that share extensive elements of a circumboreal arctic-alpine flora, the Sierra Ne- vada has developed a unique component to its alpine flora under the influence of mediterranean- climate conditions with relatively dry summers added to other alpine environmental factors of stress. Also significant in the evolution of this alpine flora has been the relative isolation of the range from other high mountain floras of the western United States. Moreover, the Sierra Nevada possesses a complex mosaic of substrate, glacial history, and soil variation superimposed over broad patterns of climatic and topographic heterogeneity. Interest in the alpine flora dates back to early descriptions by Coville (1893) and Harshberger (1911), who recognized the distinctiveness of the Sierran alpine flora. Hall and Grinnell (1919) gave a very brief description of the alpine zone in the context of a broader description of California life zones, and provided a short list of character- istic species. More significant, however, have been five studies over the past 80 years that have provided an analysis of the diversity and floristic affinities of the high elevation flora of the Sierra Nevada. The earliest of these was the work of Smiley (1921), whose definition of the boreal region of the Sierra Nevada comprised the Canadian, Hudsonian, and Arctic-Alpine zones as characterized in the Merriam system of life zones (Daubenmire 1938). These life zones roughly correspond to the upper montane, subalpine and alpine zones under current con- cepts (Fites-Kaufman et al. 2007). Smiley’s work was followed by the classic investigation of Sharsmith (1940), and in more recent decades with analyses by Chabot and Billings (1972), Major and Taylor (1977), and Stebbins (1982). Early speculations on the origin of the Sierran alpine flora were contributed by Went (1948, 1953). Beyond these broad floristic surveys, there have been numerous studies of the floristics and vegetation of regional areas of subalpine and alpine vegetation in the Sierra Nevada (Howell 1944, 1951; Klikoff 1965; Pemble 1970; Taylor 1976b; Major and Taylor 1977; Tatum 1979; Benedict and Major 1982; Burke 1982; Ratliff 1982: Benedict 1983; Porter 1983: Constantine- Shull 2000; Sawyer and Keeler-Wolf 2007). None of the existing literature has provided a satisfactory answer to the fundamental question. How many species are there in the alpine flora of 154 MADRONO the Sierra Nevada? The objective of this paper is to present a broad overview of the alpine flora of the Sierra Nevada by providing a detailed and updated analysis of the floristic richness, ecolog- ical diversity, and biogeographic relationships of the species present within the alpine zone. The paper takes a conservative approach following Sharsmith (1940) by defining the alpine zone using a lower elevational limit of 3500 m. Climatic treeline typically occurs from 3300-— 3500 m in the central and southern Sierra Nevada where the great majority of alpine habitat in California is located (Fig. 1). Although the northern Sierra Nevada lacks high elevation areas, it nevertheless has a good representation of alpine species that reach above 3500 m in the central or southern areas of the range. To provide a broader context examining the significance of elevation in the definition of the alpine zone, analyses have been made for all species occurring at or above 3300 m within California. Beyond an intrinsic interest in the evolution of biodiversity of alpine biota, there are very significant reasons to support Sierran alpine studies that can serve as baseline studies for important early warning systems of potential environmental impacts of climate change. Cli- mate change models for California suggest that there will be significant effects on environmental conditions of subalpine and alpine habitats of the Sierra Nevada (Hayoe et al. 2004; Shafer et al. 2001), and historical data on vertebrate distribu- tion demonstrates that these effects are ongoing today in influencing the distributions of verte- brate species (Moritz et al. 2008; Tingley et al. 2009). MATERIALS AND METHODS The Jepson Manual, 2nd Edition (Baldwin et al. 2012) was used to identify California species with an elevational distribution up to or above 3300 m within the state, and which occurred in the Sierra Nevada. This reference is the sole source and reference for binomials used in this article. Species at or above 3500 m in California were considered to comprise the alpine flora. The upper and lower elevational ranges of each of these species were recorded, along with their biogeographic distribution and occurrence within the geographic regions of California (Hickman 1993). These geographic regions included records of species presence in the montane and higher elevations of the northern, central, and southern subregions of the Sierra Nevada, as well as the high Cascade Range, the Klamath/Siskiyou mountains, Transverse and Peninsular ranges of southern California, and ranges east of the Sierra Nevada including the Sweetwater and White- Inyo mountains (Fig. 1). The elevational limits and geographical ranges listed in Baldwin et al. [Vol. 58 (2012) are specimen-based records and thus considered reliable. Only native species were included in this analysis, however, alien species recorded as occurring at high elevations in the Sierra Nevada are very few. Poa pratensis L. is recorded as reaching 3500 m and Taraxacum officinale F. H. Wigg. reaches 3300 m. Each taxon occurring at elevations of 3300 m or above was categorized into a series of growth forms, based on a modified scheme of Raunkiaer (1934). These categories were broad-leaved her- baceous perennials (tussocks, rosettes, and bien- nials), graminoid perennials, mats and cushion plants, geophytes, aquatics, annuals, subshrubs, woody shrubs (deciduous and evergreen), and trees. The lower elevational limit of occurrence in California was used to separate alpine species into categories of lowest elevational zone of occur- rence on the following basis: 1) foothill habitats of woodland and chaparral— <1199 m; 2) lower montane habitats dominated by mixed conifer and yellow pine forests— 1200-1999 m; 3) upper montane habitats of red fir and lodgepole pine forests— 2000-2699 m; 4) subalpine habitats of open conifer stands near treeline— 2700-3499 m; and 5) alpine habitats— >3500 m. Because elevational boundaries of these major vegetation zones change with latitude, as well as locally with slope exposure, these elevational ranges represent averaged boundaries across the west slope of the central and southern Sierra Nevada. The biogeographic range of each alpine species was classified into one of six categories. These were: 1) widespread species present in many habitats or regions across North America and/or throughout the world; 2) cordilleran species broadly distributed in mountain regions of the western United States; 3) Sierra/Cascade species with a Pacific Northwest distribution; 4) Intermountain Region species present in the Great Basin; 5) species endemic to the Sierra Nevada; and 6) species endemic to California, broadly defined to include adjacent Great Basin ranges extending into western Nevada (_.e., Sweetwater, Wassuk, and White-Inyo moun- tains) and southern Oregon. Dividing species into such simple biogeographic categories is inherently arbitrary for some species, and expanded field studies in the future may well change these classifications and alter the list of Sierran endemics based on new records or taxonomic revisions. RESULTS The Geography of California Alpine Habitats The elevational contour interval of 3500 m is highly irregular in the Sierra Nevada, as it defines a relatively continuous area along the crest of the 2011] RUNDEL: SIERRA NEVADA ALPINE FLORA iS) FiG. 1. Topographic map of California showing the major areas of mountain systems. W/I = White-Inyo Mountains, T = Tehachapi Mountains, SG = San Gabriel Mountains, SB = San Bernardino Mountains, and P = Peninsular Ranges. central and southern crest of the range extending from northern Tuolumne and Mono counties in the area of Leavitt Peak (3527 m) near Sonoran Pass and south across Yosemite National Park where the highest peak is Mount Lyell (3999 m; Fig. 2). Further south this belt of alpine habitat continues into Kings Canyon and Sequoia National parks where there are extensive areas of alpine habitat with ten peaks that reach above 4000 m. Mount Whitney at 4421 m is the highest 156 MADRONO [Vol. 58 } * . é “ a SW BETWATER T TN Dy 7 ~~ me 38° of ite / 7s 7s ’ . ’ ‘ + fi < ’ WASSUK & ¢ Pd . +94) ’ hy ~ 120 P THOGA PASS ? ca , ¢ YOSEMITE , ~ br - @ MA YE} - tal ELL ¢ a ‘ fa ‘ _ * MAMMOTH PASS ’ *. #F ¢ ‘ ’ ’ a ‘ A a ’ ¢ ad ~ 4 : 7 mae sad Py KAISER RIDGE ~ \ Rs ’ ¢ = _ ’ ‘ 37 6 ’ ~ & a ~ . . *» ’ ‘ ’ KINGS CANYC iN 4 ¢ . ¢ = a e . ’ 7 e ~ . ‘ “4 , bear ’ yA 6) co: 60 75 ‘100 8 . ‘ ’ ee ees eee Meee “sequoia cmimenie: kiometers Py J “4 ¢ ’ 4 (e) ‘ U OWENS LAKE ’ ~ ’ - fe ‘ wey . yg OLANCHA PEAK. Me ta FiG. 2. Geographic distribution of high elevations of the Sierra Nevada and adjacent ranges. The solid line marks a rough position of the 3000 m contour. Adapted from Hovanitz (1940). point in the contiguous United States. The end of the continuous chain of glaciated peaks in southern limit of this extensive and virtually the Sierra Nevada. To the south, the alpine zone contiguous alpine zone occurs at Cirque Peak reappears on Olancha Peak (3698 m; Fig. 2), the (3932 m) in Sequoia National Park at the southern southernmost glaciated summit of the range lying 2011] on the Tulare-Inyo county line (Howell 1951; Tatum 1979). Two major breaks with subalpine elevations but not true alpine provide the only major discontinuity for this primary Sierran alpine region. These are Tioga Pass in Yosemite National Park (3031 m) and Mammoth Pass (Muinaret Summit) (2824 m), which is the route for California Highway 203. The crest of the Sierra Nevada lies at lower elevations to the north of the Tioga Pass area, with only scattered areas of good alpine habitat present. A notable ecological change occurs north of this pass where volcanic substrates replace the granites of the central and southern Sierra Nevada. Fragmented communities of alpine species are present at elevations below 3500 m, particularly along exposed ridgelines and on steep north-facing slopes that were heavily glaciated. However, there are no eleva- tions in the northern Sierra Nevada that reach the 3500 m limit used here to define the alpine zone. Alpine habitats are weakly developed in Alpine Co. (with Sonora Peak reaching 3493 m) and eastern El Dorado Co. (with Freel Peak reaching 3318 m), extending to their northern limit on Mount Rose (3285 m) in the Carson Range east of Lake Tahoe in Nevada (Fig. 2). Nevertheless, there are scattered communities of alpine-like habitat existing at upper elevations in the northern Sierra Nevada, positioned above local edaphically-controlled treelines, and the alpine flora is well represented (Smiley 1915). Despite the floristic relationships of high eleva- tion Sierran species all along the range, Stebbins and Major (1965) linked the Sierra Nevada north of Lake Tahoe with the Cascade Range rather than with the region of the central and southern Sierra Nevada on the basis of the dominance of volcanic substrates. To the north of the Sierra Nevada, Mount Shasta in the southern Cascade Range reaches an elevation of 4322 m, while Lassen Peak is lower at 3187 m. The highest peaks in the Klamath Mountains of northwestern California and adjacent Oregon are Mount Eddy (2750 m) in Siskiyou Co., Thompson Peak (2744 m) in Trinity Co., and Mount Ashland (2296 m) in Jackson Co., Oregon. These high peaks contain areas with permanent or long-lasting snowfields on north-facing slopes with associated alpine species (Howell 1944; Major and Taylor 1977). There are several high mountain ranges to the east of the Sierra Nevada at the western margin of the Great Basin. The Sweetwater Mountains, located just 33 km east of the Sierra Nevada, reach 3552 m on Mount Patterson (Hunter and Johnson 1983). The Wassuk Range in west- central Nevada lie 48 km east of the Sweetwater Mountains and 88 km north of the White Mountains, reaching 3427 m on Mount Grant (Bell and Johnson 1980). The White Mountains RUNDEL: SIERRA NEVADA ALPINE FLORA 157 have an extensive alpine area and reach to 4344 m on White Mountain Peak, the third highest peak in California (Rundel et al. 2008). To the south, Mount Waucoba forms the high point at 3390 in the Inyo Mountains. The Panamint Mountains lying east of the White-Inyo Mountains reach a maximum elevation of 3366 m on Telescope Peak. Further south, the Spring Mountains in southwestern Nevada divide the Pahrump Valley and Amargosa River basins from the Las Vegas Valley watershed and define part of the south- western boundary of the Great Basin. The highest point is Charleston Peak at 3633 m. High elevations are also present in the Trans- verse and Peninsular ranges of southern Califor- nia (Fig. 1) where a subset of Sierran alpine species 1s present in weakly developed alpine-like communities (Hall 1902; Parish 1917; Horton 1960; Hanes 1976; Major and Taylor 1977; Meyers 1978). Mount San Gorgonio in the San Bernardino Mountains reaches 3506 m, while other high points are Mount San Jacinto in the San Jacinto Mountains at 3302 m and Mount Baldy in the San Gabriel Mountains at 3068 m. Alpine species are present in both xeric and mesic habitats at high elevation, but alpine communi- ties, defined as extended areas dominated by assemblages of alpine species, are only poorly developed. The alpine zone of the Sierra Nevada experi- ences mediterranean-type climate conditions that differ significantly from those that characterize the Rocky Mountains and most of the continental alpine habitats of the world where summer rainfall predominates. The fraction of annual precipitation that falls as winter snow in the Sierra Nevada is about 95% at upper treeline (Stephen- son 1998). Deep snow packs and cool temperature at higher elevations mean that snowmelt extends into the spring, but the length and magnitude of the summer drought period experienced by plants is significant. Patterns of rainfall decline gradually from north to south in the Sierra Nevada, and summer drought decreases as elevation increases because of both increased levels of precipitation and cooler temperatures with lower evaporative demand at higher elevations (Stephenson 1998; Urban et al. 2000). Winter mean monthly low temperatures are moderate in the Sierra Nevada compared to the Rocky Mountains, and soils only rarely freeze to even moderate depth. While, the mean minimum temperature above treeline is below freezing for ten months of the year, with nighttime lows that typically reach only —3 to —6°C, although extremes can reach temperatures of —15°C or lower on the high peaks. Nevertheless, these moderate low temperatures as well as other limiting factors for survival at high elevations sharply reduce the diversity of species able to tolerate such conditions (Korner 2003). 158 MADRONO Number of species/taxa 3100 3300 3500 3700 [Vol. 58 » Taxa WB Species 3900 4100 4300 4500 Elevation (m) Fic. 3. Floristic Richness The alpine flora of the Sierra Nevada, defined as species reaching 3500 m or more at their upper limit of distribution, comprises 385 vascular plant species. The species total includes 10 ferns and fern relatives (2.6%), five conifers (1.3%), 85 monocots (22.1%), and 285 eudicots (74.0%). With the inclusion of an additional 24 named varieties and subspecies, the total number of alpine taxa is 409. Of course, the predetermined elevational boundary has a very strong influence on the size of the flora (Fig. 3). If the alpine flora were defined as those species reaching to 3400 m, then 76 additional species would be added for a total of 460 species (488 taxa). Were the limit defined as 3300 m, there would be a flora of 536 species (570 taxa), with the relative proportions of monocots and eudicots virtually unchanged and the addition of five ferns and one conifer. There are 97 species (101 taxa) with an elevational range that extends as high as 4000 m, an elevation reached by only the highest Sierran peaks (Fig. 3). This number declines to 27 species that reach 4200 m in elevation. These 27 high elevation species do not display dominance by a few families as is the case with the full alpine flora but are rather spread among 15 different families (Appendix 1). Three species have been recorded as reaching to 4400 m. These are Epilobium anagallidifolium Lam. (Onagraceae), Saxifraga hyperborea R. Br. (Saxifragaceae), and Erigeron vagus Payson (Asteraceae). Additional taxa that occur up to or above 4300 m are Erigeron compositus Pursh (Asteraceae), Boechera lemmo- nii (S. Watson) W. A. Weber (Brassicaceae), Cerastium beeringianum Cham. & Schltdl. (Car- yophyllaceae), Calyptridium umbellatum (Torr.) Greene (Montiaceae), Festuca brachyphylla Schult. & Schult. subsp. breviculmis Fred., Poa keckii Soreng. and P. lettermannii Vasey (Poa- Elevational distribution of species in the high mountain flora of the Sierra Nevada. ceae), Phlox pulvinata (Wherry) Cronquist (Po- lemoniaceae), Ranunculus eschscholtzii Schltdl. var. oxynotus (A. Gray) Jeps. (Ranunculaceae), and Potentilla pseudosericea Rydb. and Sorbus californica Greene (Rosaceae). There are six families that contribute 20 or more taxa to the alpine flora. The largest of these is the Asteraceae with 55 species (59 taxa), followed in size by the Poaceae (39 species, 47 taxa), Brassicaceae (34 species), Cyperaceae (31 species), Rosaceae (21 species, 23 taxa), and Fabaceae (18 species, 27 taxa). These six families together comprise 52% of the alpine flora. At the generic level, Carex stands out prom- inently with 29 species in the alpine flora, with an additional 18 species present at elevations be- tween 3300 and 3500 m. Next in order of size are Draba (Brassicaceae, 14 species), and Lupinus (Fabaceae, 11 species, 16 taxa). There are 10 species of Boechera (Brassicaceae) and nine species each of Epilobium (Onagraceae), Eriogo- num (Polygonaceae), and Potentilla (Rosaceae). There are three genera with eight species— Penstemon (Plantaginaceae), Poa (Poaceae), and Salix (Salicaceae). Growth Form Distribution Herbaceous perennial growth forms, broadly defined, comprise the great majority of taxa reaching to or above 3500 m in the Sierra | Nevada. This growth form with all of its | subgroups includes 343 taxa, or 84% of the 409 | taxa that comprise the flora. These herbaceous | perennials can be broken down into subgroups of erect herbaceous perennials, perennial grami- noids, prostrate mats and cushion plants, bienni- | als, and geophytes. The largest numbers of | herbaceous perennials form the category of erect , herbaceous perennials, with 186 species (206 taxa; Fig. 4). The most important families for the erect , herbaceous perennials are the Asteraceae, Brassi- | 2011] 200 160 120 80 Number of taxa erect mat / annual perennial cushion graminoid geophyte RUNDEL: SIERRA NEVADA ALPINE FLORA lest) aquatic shrub suburb tree Fic. 4. Growth form distribution of alpine taxa in the alpine flora of the Sierra Nevada. caceae, Fabaceae, Rosaceae, Polygonaceae, and Onagraceae. Among the erect herbaceous peren- nials are four species that are reported to have the potential to survive as facultative annuals. Although these have not been studies in detail, it is expected that these species have biennial or short-lived perennial life histories in the alpine zone, and they are included here in the totals for erect herbaceous perennials. Three of these are members of the Brassicaceae, each representing a different genus. Among these facultative annuals, only Androsace septentrionalis L. (Primulaceae) with an elevation range of 2700-3600 m can be considered as a subalpine and alpine specialist. There are several additional groups classified broadly as erect herbaceous perennials. The 10 species of ferns and fern relatives included here within the alpine flora represent four families (Pteridaceae, Ophioglossaceae, Woodsiaceae, and Selaginellaceae). Four of these species reach elevation at or above 4000 m—Botrychium lineare W. H. Wagner, B. paradoxum W. H. Wagner, Cystopteris fragilis (L.) Bernh., and Selaginella watsonii Underw. Lowering the characterization of the alpine zone lower limit to 3300 m would add five additional fern species (Appendix 2). Also classified as erect perennials are seven species of hemiparasites, all members of the Orobanchaceae, with four species (five taxa) of Castilleja and three species of Pedicularis. Six more species from this family would be added by lowering the alpine boundary to 3300 m, includ- ing five more species of Castilleja. Next in diversity among the herbaceous perennials is the subgroup of graminoids (Cyper- aceae, Juncaceae, Juncaginaceae, and Poaceae) with 83 species (85 taxa, Fig. 4). All of the members of these four families within the alpine flora are perennials, with Agrostis, Bromus, Carex, Elymus, Juncus, Luzula, Poa, and Stipa forming genera with five or more taxa (Appendix 1). These perennial graminoids include one species of Cy grass, Muhlenbergia richardsonis Rydb. (Sage and Sage 2002). Two other C, members of this genus, the perennial M. montana Hitche. and the annual M. filiformis Rydb., just miss inclusion, reaching to elevations of 3420 m and 3350 m, respectively. Lowering the alpine boundary to 3300 m would add significantly to the diversity of graminoid perennials, with 43 additional taxa present (Appendix 2). Prostrate mats and cushion forms of growth are common in some of the herbaceous perenni- als of the Sierran alpine flora (Fig. 4). These species are low in stature and form a heteroge- neous group that shares the characteristic of a prostrate growth form with either a central taproot or multiple points of rooting through layering. Mats and cushions often form an ecologically significant component of plant cover on exposed ridges and fellfield. There are 46 species classified here as mats or cushions, with 19 of these high subalpine and alpine specialists not occurring below 2700 m elevation The growth form characteristics of mats and cushions may be genetic in some cases but in others is environ- mentally induced, with mat forms of growth only occurring at higher elevations (personal observa- tions). Alpine mat and cushion species are well represented in the Asteraceae with 13 species (notably taxa of Antennaria and Erigeron), Polygonaceae (Eriogonum) with eight species, 160 MADRONO 50 x + os 30 O - ¢ & OC 20 E A oh 2 AA 10 0 [Vol. 58 @ annuals Cishrubs a subshrubs > ® cushion/mat 3100 3300 3500 3700 3900 4100 4300 4500 Elevation (m) FIG. 5. Sierra Nevada. Brassicaceae (Draba and Anelsonia) with s1x species, and Fabaceae (Astragalus, Lupinus, Oxytropis, and Trifolium) with five species. Also notable in their ecological dominance are mats of Caryophyllaceae (Cerastium, Eremogone, Min- uartia) and Polemoniaceae (Ph/ox). The alpine flora includes just four species of geophytes, which represent the Alliaceae (A//ium obtusum Lemmon var. obtusum), Liliaceae (Ca- lochortus leichtlinii Hook. f.), Melanthiaceae (Veratrum californicum Durand var. californi- cum), and Themidaceae (Triteleia dudleyi Hoo- ver). The highest elevation species among these 1s C. leichtlinii, which reaches up to 4000 m. None of these geophytes can be considered to be high elevation specialists as all reach lower elevational limits of 1200—1500 m in California. There are six species of geophytes that just miss reaching the lower alpine limit, as defined here, but occur at or above 3300 m (Appendix 2). These include A//ium validum S. Watson (Alliaceae), /ris missouriensis Nutt. (Iridaceae), Lilium kelleyanum Lemmon (Liliaceae) and three Orchidaceae (Platanthera dilatata (Pursh) Lindl. ex L. C. Beck var. leucostachys (Lind.) Luer, P. sparsiflora Schlitr., and Spiranthes romanzoffiana Cham. Only a single species of aquatic plant, Calli- triche palustris L. (Plantaginaceae), reaches the alpine zone of the Sierra Nevada. This is perhaps not surprising given the relatively small area of oligotrophic lakes that are present above 3500 m. Potamogeton robbinsii Oakes (Potamogetona- ceae) and Limosella acaulis Sessé & Moc. Elevational distribution of taxa of annuals, shrubs, subshrubs, and mats/cushions in the alpine flora of the (Scrophulariaceaeae) have a range that extends as high as 3300 m, and a number of aquatic species including /soetes (Isoetaceae) reach eleva- tions of 3000 m. Plants with an annual life history comprise a small but significant component of the alpine flora of the Sierra Nevada, with 24 species (26 taxa) reaching to elevations of 3500 m (Fig. 4). The annual species occurring at the highest elevation is Gayophytum decipiens F. H. Lewis & Szweyk. (Onagraceae), which ranges up to 4200 m. Five other species of annuals reach 4000 m in elevation—Gentianopsis holopetala (A. Gray) Iltis (Gentianaceae), Phacelia hastata Douglas ex Lehm. subsp. compacta (Brand) Heckard (Boraginaceae), Mimulus suksdorfii A. Gray (Phrymaceae), Gayophytum racemosum Torr. & A. Gray (Onagraceae), and Collinsia torreyi A. Gray var. wrightii (S. Watson) I. M. Johnst. (Plantaginaceae). The number of annual species present increases sharply below the limit set here for inclusion in the alpine flora. Including the above taxa, there are 33 annual species (36 taxa) with a range reaching to or above 3400 m and 38 species (41 taxa) occurring at or above 3300 m (Fig. 5). Most of the annuals reaching into the alpine zone are species with wide elevational ranges that extend down to lower foothill habitats. Only 13 of the alpine annual species have ranges limited to elevations at or above 1200 m, a distribution that would indicate adaptation to montane and higher elevation habitats. Five annual taxa can be 2011] considered as subalpine and alpine specialists having a lower elevation limit of 2700 m or above and/or a median elevational range above 3000 m. These species, none of which ranges as high as 4000 m or above, are Comastoma_ tenellum (Rottb.) Toyok. (Gentianaceae), Cryptantha cir- cumscissa (Hook. & Arn.) I. M. Johnst.var. rosulata J. T. Howell (Boraginaceae), Strep- tanthus gracilis Eastw. (Brassicaceae), and Lepto- siphon oblanceolatus (Brand) J. M. Porter & L. A. Johnson and Gymnosteris parvula A. Heller (Polemoniaceae). Just missing this criteria, but certainly also a high elevation specialist, is Phacelia orogenes Brand (Boraginaceae). Four of these six, with Comastoma tenellum and Gymnosteris parvula as exceptions, are Sierra Nevada endemics. The most important family in contributing to the annual flora of high elevations is the Boraginaceae, with 11 species (12 taxa) repre- senting five genera. Next in importance are the Polemoniaceae with five species (comprising five genera), and the Onagraceae with four species (five taxa) representing just a single genus. There are four genera that contribute three or more species to the annual flora. These are Gayophy- tum (Onagraceae, four species, five taxa), Phace- lia (Boraginaceae, four species), Cryptantha (Boraginaceae, three species, four taxa), and Mimulus (Phrymaceae, three species). Subshrubs, defined as semi-woody species that maintain living perennial tissue in winter above the ground surface, include 13 species occurring at elevations of 3500 m or above (Fig. 4). The Asteraceae contribute more than 60% of the alpine flora of subshrubs, with eight species. Four species of Ericameria (Asteraceae) and three species each of Penstemon (Plantaginaceae), and one Monardella (Lamiaceae) form subshrubs that reach alpine elevations. Five species are consid- ered to be subalpine and alpine specialists based on a lower elevational limit of 2700 m or a mean elevational range above 3000 m. Four of these are members of the Asteraceae—Sphaeromeria cana (D. C. Eaton) A. Heller, Ericameria parryi (A. Gray) G. L. Nesom & G. I. Baird var. monocephala (A. Nelson & P. B. Kenn.) G. L. Nesom & G. I Baird, E. bloomer (A. gray) J. F. Macbr., and Chrysothamnus viscidiflorus (Hook.) Nutt. var. viscidiflorus. The two latter species have very broad elevational occurrence from 800-4000 m. There are 23 species of woody shrubs that extend into the alpine zone of the Sierra Nevada (Fig. 4). Just four families account for the majority of the high elevation shrubs. The largest of these is the Salicaceae (eight species of Salix), followed by the Ericaceae (five species, each in a different genus), Grossulariaceae (three species of Ribes), and Rosaceae (three species, each in a different genus). The highest elevation reached is RUNDEL: SIERRA NEVADA ALPINE FLORA 16] reported for Sorbus californica at 4300 m. How- ever, this elevation record appears to not be supported by specimen records in the Consortium of California Herbaria (ucjeps.berkeley.edu/ consortium), and therefore needs confirmation. There are six additional shrub species that reach elevations of 4000 m—Salix orestera C. K. Schneid., S. planifolia Pursh, S. petrophila Rydb., Gaultheria humifusa (Graham) Rydb., Holodiscus discolor (Pursh) Maxim. var. microphyllus (Rydb) Jeps., and Ribes montigenum McClatchie. Only three shrub species can be considered as subal- pine and alpine specialists based on a lower elevational limit at or above 2700 m or median range of occurrence above 3000 m. These are Salix planifolia, S. brachycarpa Nutt. var. bra- chycarpa, and S. nivalis Hook. Three more shrub species just miss this definition of high elevation specialist. Arctostaphylos uva-ursi (L.) Spreng. has an elevational range of 2400-3300 m, while Jamesia americana Torr. & A. Gray (Hydrangea- ceae) and Ribes cereum Douglas var. inebrians (Lindl.) C. L. Hitche. are alpine species that extend down to lower elevations of 2070 m and 2100 m, respectively. Including the above species, there are a total of 39 shrub species that occur at elevations of 3300 m or above in the Sierra Nevada. This group includes two more species of Salix, one additional Ribes, five Ericaceae, three Rosaceae, two species of Caprifoliaceae, and a scattered diversity of species from other families (Appendix 2). Five species of coniferous trees in the Pinaceae have scattered populations that extend well above typical treeline elevation on favorable sites. The treeline pines, Pinus albicaulis Engelm., P. flexilis E. James and P. balfouriana Grev. & Balf., all have local populations that reach as high as 3700 m in elevation in the Sierra Nevada, while P. contorta Loudon subsp. murrayana (Grev. & Balf.) Critchf. and Tsuga mertensiana (Bong.) Carriere reach 3500 m. Just missing the elevation of the alpine zone are scattered trees of Pinus monticola Douglas ex D. Don that reach up to 3400 m. Elevational Amplitude Separating alpine taxa into categories of elevational ranges over which they occur provides some insight into their ecological amplitude and thus a crude measure of potential niche breadth. There are nine obligate alpine taxa in the Sierra Nevada restricted in occurrence to elevations at or above 3500 m. These are Boechera depaupe- rata (A. Nelson & P. B. Kenn.) Windham & Al-Shehbaz (Brassicaceae), Botrychium para- doxum and B. tunux Stensvold & Farrar (Ophio- glossaceae), Carex incurviformis Mack. (Cyper- aceae), Draba sierra Sharsm. (Brassicaceae), Eriogonum wrightii Torr. ex Benth. var. olanchense 162 140 120 100 80 60 Number of taxa 40 20 foothill lower montane FIG. 6. MADRONO upper montane [Vol. 58 subalpine alpine Lower elevational zone of occurrence for taxa in the alpine flora of the Sierra Nevada. The elevational limits of these zones are 0—1190 m for the foothill zone, 1200-1990 m for the lower montane zone, 2000—2699 m for the upper montane zone, 2700-3490 m for the subalpine zone, and 3500 m and above for the alpine zone. (J. T. Howell) Reveal (Polygonaceae), Minuartia stricta (Sw.) Hiern (Caryophyllaceae), Phlox dispersa Sharsm. (Polemoniaceae), and Poa let- termanii (Poaceae). There an additional 67 alpine species (70 taxa, 17.2% of all taxa) with ranges restricted to the elevations of subalpine and alpine habitats at or above 2700 m (Fig. 6). Examining the floristic composition of all 76 species (79 taxa) with a range restricted to subalpine and alpine eleva- tions, just four families comprise more than half of these. These are the Asteraceae (11 species), Brassicaceae (nine species), Rosaceae (eight species, nine taxa), and Poaceae (eight species). Three additional species would be added to the obligate subalpine and alpine flora if the eleva- tional limit were reduced to 3300 m. These are Astragalus ravenii Barneby (Fabaceae), Carex tiogana D. W. Taylor & J. D. Mastrog. (Cyper- aceae), and Chaenactis douglasii (Hook.) Hook. & Arn. var. alpina A. Gray (Asteraceae). Looking at the level of all alpine taxa, 22.4% have a lower elevational limit in the upper montane zone (2000-2699 m) and a further 31.0% have a lower limit in the lower montane zone (1200-1999 m). Finally 27.5% of the alpine taxa have a broad elevational amplitude of occurrence extending upward from the foothill zone below 1200 m up into the alpine (Fig. 6). Plotting the elevational amplitude of all of the alpine taxa shows a peak at about 2300 m, with relatively fewer species exhibiting very broad or very narrow elevational amplitudes (Fig. 7). Nevertheless, there are many taxa with surprising broad ranges of elevational occurrence. There are 77 species that have an elevational amplitude of 3000 m or more, and six species that have 4000 m or more of amplitude in California. These latter species, each in a different family, are Callitriche palustris (Plantaginaceae), Calyptridium umbella- tum (Montiaceae), Cystopteris fragilis (Woodsia- ceae), Draba cana Rydb. (Brassicaceae), Epilo- bium ciliatum Raf. subsp. ciliatum (Onagraceae), and Erysimum capitatum (Hook.) Greene var. capitatum (Brassicaceae). Were the elevational definition of the alpine zone lowered to 3300 m, a large number of species with broad elevational amplitudes would be added to the flora. There are 42 species in this group of added taxa that have 3000 m or more of elevational amplitude in their range of occurrence. Biogeography and Endemism Within the Sierra Nevada itself, the distribu- tions of the high elevation flora are relatively well spread between the northern, central and south- ern subregions of the Sierra Nevada. Assessing species reaching an elevational boundary of 3300 m, 70% of the 567 taxa occur in all three subregions. The northern subregion has 76% of the alpine flora present, while the central and southern Sierra Nevada have 90% and 88% of the alpine flora present, respectively. A number of alpine species have their southern limit of distribution in the central Sierra Nevada. These include Carex whitneyi Olney (Cyperaceae), Podistera nevadensis (A. Gray) S. Watson (Apia- ceae), Claytonia megarhiza (A. Gray) S. Waston (Montiaceae), Thalictrum alpinum L. (Ranuncu- laceae), Galium grayanum Ehrend. var. grayanum (Rubiaceae), and Salix nivalis (Salicaceae). | 2011] RUNDEL: SIERRA NEVADA ALPINE FLORA 163 60 50 40 fae) & xe) 5 30 Q = =) Zz 20 10 0 0 400 800 1200 1600 2100 2400 2800 3200 3600 4000 4400 Elevational amplitude (m) FIG. 7. Elevational amplitude of alpine taxa in the Sierra Nevada. These values are based on the range expressed from upper and lower limits of elevational distribution in California as presented in Baldwin et al. (2012). The broader biogeographic relationships of the alpine flora at or above 3500 m indicate its diverse origins (Fig. 8). Widespread species dis- tributed across North America and beyond as boreal or arctic-alpine taxa comprise 13.6% of the flora (Table 1). The largest group of taxa (34.3%) shows patterns of distribution as cordil- leran species widespread in mountain regions of the western United States. Next in importance 160 140 120 100 Number of taxa cord S-C wide FIG. 8. are taxa with a range in the Intermountain Region of the Great Basin, comprising 20.5% of taxa. A group consisting of 15.8% of the taxa has ranges extending along the Sierra Nevada axis to the Cascade Range and often on to the Pacific Northwest. The alpine flora of the Sierra Nevada includes 36 endemic taxa restricted in their distribution to the Sierra Nevada (Table 2). These endemic taxa end end-cal int Biogeographic relationships of the alpine flora of the Sierra Nevada. WIDE = widespread taxa present in many habitats or regions across North America and/or throughout the world; CORD = cordilleran taxa widespread in mountain regions of the western United States; S-C = Sierra/Cascade taxa with a Pacific Northwest distribution; END = taxa endemic to the Sierra Nevada; and END-CAL = taxa endemic to California, as broadly defined; INT = intermountain taxa present in the Great Basin. See text for discussion. 164 TABLE 1. SIERRA NEVADA. Biogeographic range category Widespread taxa distributed across North America and beyond as circumboreal or arctic-alpine taxa MADRONO [Vol. 58 EXAMPLES OF THE BROADER BIOGEOGRAPHIC RELATIONSHIPS OF THE ALPINE FLORA OF THE Examplar taxa Anemone drummondii (Ranunculaceae), Carex capitata (Cyperaceae), Crepis nana (Asteraceae), Cystopteris fragilis (Woodsiaceae), Deschampsia cespitosa (Poaceae), Oxyria digyna (Polygonaceae), Phleum alpinum (Poaceae), Rhodiola integrifolium (Crassulaceae), Salix nivalis and S. petrophila (Salicaceae), Sibbaldia procumbens (Rosaceae), Trisetum spicatum (Poaceae) Cordilleran taxa widespread in mountain regions of the western United States Antennaria media (Asteraceae), Carex phaeocephala (Cyperaceae), Erigeron vagus (Asteraceae), Gentiana calycosa (Gentianaceae). Lewisia pygmaea (Montiaceae), Phlox condensata (Polemoniaceae), Poa glauca subsp. rupicola (Poaceae), Ribes cereum (Grossulariaceae), Stipa pinetorum (Poaceae) Intermountain Region taxa distributed across the Great Basin Cryptantha nubigena (Boraginaceae), Cymopterus cinerarius (Apiaceae), Eriogonum incanum (Polygonaceae), Ivesia shockleyi (Rosaceae), Jamesia americana (Hydrangeaceae), Podistera nevadensis (Apiaceae), Selaginella watsonii (Selaginellaceae), Trifolium monanthum subsp. monanthum (Fabaceae) Taxa extending from the Pacific Northwest and Cascade Range Anelsonia eurycarpa (Brassicaceae), Astragalus whitneyi (Fabaceae), Boechera howellii (Brassicaceae), Carex breweri (Cyperaceae), Eriogonum lobbii (Polygonaceae), Gentiana newberryi (Gentianaceae), Potentilla breweri and P. flabellifolia (Rosaceae), Ranunculus alismifolius var. alismellus (Ranunculaceae), Senecio fremontii var.occidentalis (Asteraceae), Silene sargentii (Caryophyllaceae) are heavily weighted toward subalpine and alpine specialists, with 29 of these restricted in distribu- tion to elevations of 2700 m or above, or with median elevational range above 3000 m. Two generic lineages are prominent among these endemics, with five taxa each of Draba and Eriogonum. Five of the alpine endemics are annual species—Orochaenactis thysanocarpha (A. Gray) Coville (Asteraceae), Cryptantha cir- cumscissa var. rosulata (Boraginaceae), Strep- tanthus gracilis (Brassicaceae), and Leptosiphon oblanceolatus (Polemoniaceae). The southern Sierra Nevada is the most significant subregion for endemics, with 15 of the 36 endemic taxa (42%) are restricted in distribution to the area from the Kings River drainage south that includes Kings Canyon and Sequoia National parks (Table 2). These are Draba cruciata Payson, D. longisquamosa O. E. Schulz, D. sharsmithii Rollins & R. A. Price, Eriogonum polypodum Small, E. spergulinum A. Gray var. pretense (S. Stokes) J. T. Howell, E. wrightii var. olanchense, Galium hypotrichium A. Gray subsp. subalpinum (Hilend & J. T. Howell) Ehrend., Leptosiphon oblanceolatus, Monardella beneolens Shevock, Ertter & Jokerst, Oreonana clementis (M. E. Jones) Jeps., Orthochaenactis thysanocarpha, Phlox dispersa, Pinus balfouriana var. austina R. J. Mastrog. & J. D. Mastrog., Streptanthus gracilis, and Trifolium kingie S. Waston subsp. dedeckerae (J. M. Gillett) D. Heller. Another 13 taxa are restricted to the central and southern Sierra Nevada. Five of the endem- ics are present across the northern, central and southern subregions of the Sierra Nevada, and three endemic taxa are restricted in occurrence to the central Sierra Nevada. These are Draba sierrae, Eriogonum ovalifolium Nutt. var. caeles- tinum Reveal, and Lupinus gracilentus Greene (Table 2). Although the absence of peaks above 3500 m in the Sierra Nevada north of Yosemite National Park explains the lack of endemics restricted to this subregion, the scattered lower elevation alpine communities of the northern Sierra Nevada retain moderately high richness of species. In addition to the members of the alpine flora that are endemic to the Sierra Nevada, there are an additional 31 alpine taxa that are Californian endemics, allowing for a broad interpretation of the floristic region to include the westernmost ranges of the Great Basin lying close to the Sierra Nevada and the southern Cascade Range in Oregon. Many of the Californian endemics have ranges that extend to the Sweetwater and/or — White Inyo mountains, while others extend into | the high Transverse and Peninsular Ranges of | southern California and a small number extend © into the southern Cascade Range. The pattern of | dominant endemism centered in the southern Sierra Nevada is not seen among these taxa. Twelve of these occur throughout the Sierra Nevada and an additional 12 are restricted to the © central and southern areas of the range, while | only four species are limited to the southern Sierra Nevada. If the lower limit of the alpine zone were | dropped to 3300 m, 11 additional Sierra Nevada endemics would be added (Table 2). Four of | these are restricted to the southern Sierra Nevada | (Astragalus ravenii, Boechera pygmaea (Rollins) | Al-Shehbaz, Castilleja praeterita Heckard & © Bacig., and Phacelia orogenes), with three each — present in the central and southern Sierra Nevada | and in all three regions. One species is restricted 2011] to the central Sierra Nevada. Five additional Californian endemics would be added if the lower alpine limit was dropped to 3300 m (Table 2). DISCUSSION Defining the Alpine Zone of California Critically defining what species should be included in an alpine flora is an imperfect task given the lack of a simple operational definition, as discussed below. The high elevation areas of the Sierra Nevada broadly classified as subalpine and alpine, or upper Hudsonian and Arctic-Alpine in the Merriam life zone classification (Daubenmire 1938), would roughly include those areas lying above about 3000 m elevation (Fig. 2). Such subalpine and alpine habitats cover extensive areas of the central and southern Sierra Nevada, but only scattered areas of the northern Sierra Nevada lying north of Sonora Pass. The higher elevation area of this northern Sierran region, however, supports mosaics of subalpine forest, shrublands, and low alpine-like vegetation (Smiley 1915, 1921). A simple definition of alpine habitat is that area occurring above treeline, with the caveat that most alpine species are not obligate in inhabiting habitats above treeline and typically occur to varying degrees at lower elevations (Packer 1974). While this approach sounds logical, timberline itself can be highly variable even in a local area depending on slope exposure, erosional history, parent material, disturbance history, and local microclimate (Billings 2000). Sharsmith (1940) recognized the alpine flora as a distinct subdivision of the overall California flora, characterized by its geographic range, growth forms, species composition, and constancy with which the alpine association of species is maintained. Although he described the alpine zone as reaching its lower limit at an average elevation of 3500 m, the limit used in this paper, nowhere in his dissertation is there a clear statement of criteria for his inclusion of species. He stated, While certain species are absolute indicators of the alpine flora, it is the particular association of species which gives it its characteristic. Although many species occur elsewhere, this special assemblage is not met until the alpine area 1s reached. Everywhere the flora presents the impression of unity, an impression rein- forced by increased field experience. The combined subalpine and alpine flora of the Sierra Nevada as defined by Smiley (1921) included 633 species, with 41 listed as indicators of the Arctic-Alpine zone. He considered 158 Species to be Sierra Nevada endemics and another 20 species to be restricted in distribution to the Sierra Nevada and the southern California mountains. Sharsmith (1940) included 189 species RUNDEL: SIERRA NEVADA ALPINE FLORA 165 in his alpine flora of the Sierra Nevada, with 31 of these considered to be endemic. This flora was composed of 183 herbaceous perennials and six annual species, but did not include any woody species. A similar estimate was made by Stebbins (1982) who stated that there were 207 species in the Sierra Nevada alpine flora. Finally, a much higher estimate came from Major and Bamberg (1967) who used the species descriptions in Munz (1959) to estimate a Sierran alpine flora of about 600 species, a number similar to that reported here for taxa reaching 3300 m or above. Growth Forms The relative dominance of a herbaceous perennial growth forms present in the alpine flora of the Sierra Nevada is typical of other alpine floras worldwide and does not change dramatically in a gradient from the Rocky Mountains west across the Great Basin (Billings 1978, 2000; Rundel et al. 2008). Herbaceous perennials have the characteristic of maintaining large proportions of total biomass belowground where they play an important role in carbohy- drate storage over the winter months (Mooney and Billings 1960; Billings 1974). The herbaceous perennials include species with a variety of ecological forms and life history strategies of carbon allocation to belowground and above- ground vegetative, and reproductive tissues (Rundel et al. 2005), and many of these are relatively long-lived plants surviving for decades (Billings 1974; Pollak 1991). As in other alpine regions, perennial grami- noids in alpine habitats of the Sierra Nevada commonly dominate plant communities of wet meadows that dry earlier than fellfield commu- nities. In contrast, fellfield habitats exhibit a mixed dominance of broad-leaved erect perenni- als, perennial graminoids, and mats and cushions (Rundel et al. 2005). Mat and cushion growth forms of herbaceous perennials are widespread in the high elevation Sierra Nevada, where the 46 taxa listed here represent 12% of the alpine flora. These are most prominent ecologically on wind- swept rocky slopes or other exposed areas that remain snow-free during the winter. Because of limiting stress factors of short and severe growing conditions, annual plants are generally rare in the typical circumboreal arctic- alpine floras of the Northern Hemisphere, comprising no more than 1—2% of the flora (Billings 2000). Although not abundant, annuals, nevertheless, are more common in alpine flora of the Sierra Nevada and White Mountains where they comprise about 6—-8% of the floras (Jackson and Bliss 1982; Jackson 1985; Rundel et al. 2008). The species richness of alpine annual taxa, however, drops rapidly at elevations above 3300 in the Sierra Nevada (Fig. 5). Went (1948, 1953) 166 MADRONO [Vol. 58 TABLE 2. ENDEMIC TAXA OF THE ALPINE FLORA OF THE SIERRA NEVADA, CALIFORNIA WITH THEIR UPPER RANGE OF OCCURRENCE AND GROWTH FORMS. For range: n = northern Sierra Nevada, c = central Sierra Nevada, and s = southern Sierra Nevada. For growth forms: A = annual, G = geophyte, P = erect herbaceous perennial, P-G = perennial graminoid, P-MAT = mat or cushion, SS = subshrub, T = tree. Species names follow Baldwin et al. (2012). Endemic group Family Range Growth form Sierra Nevada endemics >3500 m Aquilegia pubescens Ranunculaceae n,c,s P Calamagrostis muiriana Poaceae C,S P-G Carex congdonii Cyperaceae C,s P-G Cryptantha circumscissa var. rosulata Boraginaceae n,Cc,s A Dodecatheon subalpinum Primulaceae C,S P Draba cruciata Brassicaceae S P Draba lemmonii Brassicaceae n,C,s P Draba longisquamosa Brassicaceae S P Draba sharsmithii Brassicaceae S P Draba sierrae Brassicaceae Cc P-MAT Eriogonum nudum var. scapigerum Polygonaceae C,S P Eriogonum ovalifolium var. caelestinum Polygonaceae Cc P-MAT Eriogonum polypodum Polygonaceae S P-MAT Eriogonum spergulinum var. pratense Polygonaceae S P Eriogonum wrightii var olanchense Polygonaceae S P-MAT Galium hypotrichium subsp. subalpinum Rubiaceae S P Hazardia whitneyi var. whitneyi Asteraceae n,c,s SS Ivesia muirti Rosaceae C,S P Ivesia pygmaea Rosaceae CS P Leptosiphon oblanceolatus Polemoniaceae S A Lewisia disepala Montiaceae c,S P Lupinus covillei Fabaceae C,S P Lupinus gracilentus Fabaceae Cc P Luzula orestera Juncaceae n,c,$S P-G Monardella beneolens Lamiaceae S SS Oreonana clementis Apiaceae S P Oreostemma peirsonii Asteraceae C,S P Orochaenactis thysanocarpha Asteraceae S A Phlox dispersa Polemoniaceae S P-MAT Pinus balfouriana var. austrina Pinaceae S T Poa stebbinsii Poaceae c,S P-G Polemonium eximium Polemoniaceae c,S P Stipa kingii Poaceae C,S P-G Streptanthus gracilis Brassicaceae S A Trichophorum clementis Cyperaceae C,S P-G Trifolium kingii subsp. dedeckerae Fabaceae s P Sierra Nevada endemics 3300-3499 m Astragalus ravenii Fabaceae S P Boechera pygmaea Brassicaceae S P Castilleja praeterita Orobanchaceae S FP Erigeron elmeri Asteraceae C,S P Hulsea vestita subsp. vestita Asteraceae C;s P Ipomopsis aggregata subsp. bridgesii Polemoniaceae C,Ss P Lilium kelleyanum Liliaceae c,s G Lomatium torreyi Apiaceae n,c,s P Phacelia eisenti Boraginaceae c,s A Phacelia orogenes Boraginaceae S A Trifolium monanthum subsp. tenerum Fabaceae n,c,s P Californian endemics >3500 Astragalus kentrophyta var. danaus Fabaceae C,S P-MAT Carex mariposana Cyperaceae n,c,s P-G Castilleja nana Orobanchaceae n,c,s P Chaenactis alpigena Asteraceae n,c,s P-MAT Delphinium polycladon Ranunculaceae n,c,s P Draba breweri Brassicaceae n,c,s P Draba subumbellata Brassicaceae S P-MAT Eriogonum gracilipes Polygonaceae C,S P-MAT 2011] TABLE 2. Endemic group Galium hypotrichium subsp. hypotrichium Hulsea vestita subsp. pygmaea Ivesia lycopodioides subsp. lycopodioides Ivesia lycopodioides subsp. scandularis Ivesia santolinoides Lewisia glandulosa Lupinus breweri var. breweri Lupinus breweri var. bryoides Lupinus latifolius var. parishii Lupinus lepidus var. ramosus Lupinus padre-crowleyi Lupinus pratensis var. pratensis Phyllodoce breweri Poa keckii Potentilla pseudosericea Potentilla wheeleri Primula suffrutescens Ranunculus eschscholtzii var. oxynotus Tonestus peirsonii Triteleia dudleyi Viola pinetorum subsp. grisea Viola purpurea subsp. mesophyta Californian endemics 3300-3499 m Eriogonum latens Frasera puberulenta Hordeum brachyantherum subsp. californicum Penstemon caesius Plagiobothrys torreyi var. diffusus suggested that many of the high elevation annuals in the Sierra Nevada were related to desert species. Severe winter conditions typically limit the occurrence of woody plants above treeline, with prostrate mats and cushions as prominent excep- tions. The upright growth form of woody shrubs and krummbholtz tree species exposes their tissues to extreme conditions of temperature and wind exposure (Korner 2003). This impact on shrub occurrence can be seen in Fig. 5 where shrub richness in the Sierra Nevada drops sharply with increasing elevation above 3300 m, similar to the pattern for annual species. Much of the alpine flora of woody species comes from species of Salix and members of the Ericaceae, groups which favor moist habitats with some level of protection. Biogeography and Endemism The alpine flora of mountain ranges on the western margin of the Great Basin of California and western Nevada exhibit very strong relation- ships to that of the Sierra Nevada (Rundel et al. 2008). The Sweetwater Mountains supports a flora of 173 species in 16 km? of alpine habitat, with 94% of this flora common to the Sierra Nevada (Hunter and Johnson 1983). The Wassuk Range has an alpine flora of 70 species in just 2.6 km’ of alpine habitat (Bell and Johnson 1980). Again, this flora is has stronger floristic RUNDEL: SIERRA NEVADA ALPINE FLORA 167 CONTINUED. Family Range Growth form Rubiaceae C,S P Asteraceae C,S P Rosaceae ee P Rosaceae C,S P Rosaceae n,C,s P Montiaceae C.S P Fabaceae n,c,s P-MAT Fabaceae S P-MAT Fabaceae C,S P Fabaceae C,S P Fabaceae S P-MAT Fabaceae Gs P Ericaceae n,c,S S Poaceae n.c.S P-G Rosaceae C,S P Rosaceae S P Primulaceae n.c.s Pr Ranunculaceae N:c.s P Asteraceae Cc Pp Themidaceae C,S G Violaceae n,c,s P Violaceae n,c,s P Polygonaceae C,S Fr Gentianaceae C,S P Poaceae n,c,S P-G Plantaginaceae S SS Boraginaceae Ie.s A relationships to the Sierra Nevada than the Rocky Mountains. As with the Sweetwater Mountains and Was- suk Ranges, the flora of the White Mountains exhibits much stronger floristic relationships to the Sierra Nevada than to the Rocky Mountains. About 90% of the species in the alpine flora of the White Mountains are also found in the Sierra Nevada (Rundel et al. 2008), compared with only 58% that occur in the ranges of the central Rocky Mountains (Scott 1995). These values are signif- icantly higher for both ranges than earlier estimates made on incomplete data (Lloyd and Mitchell 1973). Mountain ranges in the central Great Basin generally show strong floristic linkages to the Rocky Mountains and weaker links to the Sierra Nevada (Billings 1978). Loope (1969) reported 189 alpine species from the Ruby Mountains in northeastern Nevada, with this flora heavily linked to the Rocky Mountains. The isolated San Francisco Mountains in Arizona with only 5.2 km’ of alpine habitat has 80 species, and likewise shows strong floristic relationships to the Rocky Mountains despite its separation of about 200 km (Schaak 1983). The level of endemism in the alpine Sierra Nevada flora is a relatively small part of the overall endemism for the montane and higher parts of the range. Based on current information, there are 205 taxa endemic to what The Jepson 168 Manual (Hickman 1993) classifies as the north- ern, central, and southern high Sierra Nevada, i.e., the montane, subalpine and alpine zones above foothill habitats (R. Moe, Univ. of California, Berkeley, personal communication). The 36 Sierran endemics present in the alpine flora would thus comprise 18% of the endemic flora of the higher Sierra Nevada. The unique California component of the alpine flora of the Sierra Nevada is considerably greater if one considers the endemic component of 31 species in the alpine flora that are not uniquely limited to the Sierra Nevada but are Californian endemics as defined earlier. Combining the endemic taxa with Sierran and Californian limits of distribution, the total of 66 taxa represents 16% of the alpine flora. This is a relatively high figure compared to other alpine ranges in continental North America and Europe, and reflects the environmental stress conditions asso- ciated with the summer-dry mediterranean-type climate present in the Sierra Nevada. Stebbins (1982) analyzed the flora of the high Sierra Nevada, defined similarly to that of Smiley (1921) as the upper montane to alpine zones, and identified 119 endemic species, 13.5% of the total flora. He further noted that another 60% of the flora extended beyond the Sierra Nevada only as far as southern California, western Nevada, and southern Oregon. Raven and Axelrod (1978) briefly discussed the diversity and evolution of the subalpine and alpine flora of the Sierra Nevada, listing 68 endemics for this region. Their table of endemics, however, is outdated by more recent information on distribution patterns and species concepts. Shevock (1996) gave a figure of 405 endemic taxa of vascular plants for the entire Sierra Nevada. The 36 alpine endemics reported here would comprise 9% of this total. Of the three geograph- ical subregions (northern, central, and southern) of the entire range, the southern Sierra Nevada is the richest in endemics, rare species, and total floristic composition (Shevock 1996), a finding similar to that reported here. The Evolution of the Sierran Alpine Flora A detailed assessment of the biogeographic and evolutionary origin of the alpine flora of the Sierra Nevada is beyond the scope of this review. Broad interpretations of biogeographic relation- ships within alpine lineages have been discussed by previous authors (e.g., Smiley 1921; Sharsmith 1940; Chabot and Billings 1972; Taylor 1977; Major and Taylor 1977; Raven and Axelrod 1978; Stebbins 1982) but recent phylogenetic studies have made many of these earlier interpre- tations subject to re-evaluation. Evidence for a north to south route of colonization of high mountain areas of the Sierra MADRONO [Vol. 58 Nevada comes from a pattern of decreasing presence of Rocky Mountain floristic elements and an increasing number of endemics alpine species as one moves from the northern to southern crest of the range (Chabot and Billings 1972; Raven and Axelrod 1978). The southern limit of a number of alpine species on Mount Lassen suggests the possibility that some of these and other Cascade Range species may well have been present in the Sierra Nevada in the late Pliocene or early Pleistocene. Although the species composition of lower and middle eleva- tion conifer forests of Lassen National Park are strongly related to that of the Sierra Nevada, the summits of the highest peaks in Lassen support an alpine flora that exhibits stronger floristic links to Mount Shasta and the Cascade Range to the north (Gillett et al. 1995). Alpine species with disjunct patterns of distribution from Mount Lassen to the Cascade Range volcanoes include Cardamine_ bellidifolia L. (Brassicaceae), Carex illota L. H. Bailey (Cyperaceae), Collomia larsenii (A. Gray) Payson (Polemoniaceae), Draba aure- ola S. Watson (Brassicaceae), Erigeron elegantu- lus Greene and E. nivalis Nutt. (Asteraceae), Hulsea nana A. Gray (Asteraceae), Polemonium pulcherrimum Hook. var. pilosum (Greenm.) Brand (Polemoniaceae), and Silene suksdorfii B. L. Rob. (Caryophyllaceae). The Klamath Moun- tains also mark the southern distribution limit of a number of boreal species that do not occur in the high elevations of the Sierra Nevada (Howell 1944). Alpine and subalpine species characteristic of wet meadows and other moist sites typically have broad geographic ranges but become increasing habitat specific moving to the south in the Sierra Nevada as precipitation decreases (Kimball et al. 2004; Moore et al. 2007). The relative isolation of the Sierra Nevada from northern ranges and the summer-dry have clearly acted as a filter to exclude some widespread circumpolar arctic- alpine species such as Dryas integrifolia Vahl (Rosaceae) and Silene acaulis L. (Caryophylla- ceae) which do not occur anywhere in California. © Species growing in xeric rocky habitats show | higher levels of endemism and smaller range size — due to isolation and divergence from ancestral | populations distributed in wetter habitats to the | north. | More controversial, however, is the origin of | disjunct Rocky Mountain species present in the — central and southern Sierra Nevada, often growing in azonal soil conditions. There is both geological and paleobotanical evidence to suggest — that the mean elevation of the Great Basin was as | much as 1500 m higher in the Miocene and that the current Basin and Range topography is the | result of subsidence rather than uplift (Wernicke © et al. 1988; Wolfe et al. 1997). The presence of | higher elevations in the Great Basin during the © 2011] Pleistocene could possibly have provided step- ping stones to allow the dispersal of alpine organisms from the east (Major and Bamberg 1967; Taylor 1976a). Molecular evidence indi- cates that at least one lineage of butterflies entered the Sierra Nevada by this route (Nice and Shapiro 2001). However, other authors feel that the majority of these disjunct plant species reached the Sierra Nevada by the same dominant route from the Western Cordillera via the Cascade Range and south (Chabot and Billings 1972). Modes of speciation in the development of the endemic alpine flora of the Sierra Nevada are clearly complex. Polyploidy and associated apo- mixis are widely recognized as major factors in plant evolution, and these factors have had a relatively recent impact on speciation in produc- ing stable self-propagating lineages (Soltis et al. 2009). In the alpine region of the Sierra Nevada, as in other alpine regions, diploid lineages of polyploid complexes often occupy unglaciated areas and resist introgression due hypothetically to a significantly higher seed set. However, asexual apomictic populations are more wide- spread than their sexual relatives in glaciated areas. Sexual and asexual polyploids may become distinct stabilized species through hybrid origin. Reproductive isolation and stability of tetra- ploids within their respective distribution as well as the value of uniparental reproduction provide the advantages of apomixis. Many important genera in the alpine flora of the Sierra Nevada are notable for the presence of apomixis, with Boechera (Schranz et al. 2005; Dobes et al. 2007), Draba (Jordon-Thaden and Koch 2008), and Antennaria (Bayer and Stebbins 1987) as examples. Additional speciose genera in the Sierra Nevada known to have complex apomictic populations include Arnica and Crepis (Aster- aceae; Noyes 2007), Poa and Calamagrostis (Poaceae), and Potentilla (Rosaceae) (Asker and Jerling 1992). Other modes of alpine speciation have also been described for the Sierra Nevada. Some speciation, for example, has hypothetically come from lowland arid-adapted taxa colonizing the glaciated terrain of the range at the end of the Pleistocene (Went 1948, 1953). Speciation has also been shown to be the result of population disjunction and reproductive isolation (Chase and Raven 1975). Although the Transverse and Peninsular rang- es are well separated from the higher elevations of the Sierra Nevada, more than one third of the Sierran alpine flora has a range of distribution that extends to these southern California ranges. While some of these species occur at lower elevations, others are typically subalpine and alpine species that must have crossed the Mojave Desert during the cold conditions of the Pleisto- RUNDEL: SIERRA NEVADA ALPINE FLORA 169 cene. This latter group includes Androsace septentrionalis (Primulaceae), Hulsea vestita A. Gray subsp. pygmaea (A. Gray) Wilken (Aster- aceae), Oxyria digyna (L.) Hill (Polygonaceae), and Podistera nevadensis (Apiaceae). There are lessons to be learned from recent studies of the patterns of diversification in the European alpine flora. These strongly demon- strate that speciation have been promoted by diverse ecological, evolutionary, and life history traits related to population structure, phyloge- netic relationships, breeding system, dispersal syndromes, ecophysiological ranges of habitat requirements, and competitive abilities (Comes and Kadereit 1998; Taberlet et al. 1998: Hewitt 2000; Gugerli and Holderegger 2001; Vargas 2003). The complex and dynamic climatic and geological history of the Sierra Nevada operating on such traits suggests that there have been a range of different colonization and extinction histories that are species specific. Much more work on the comparative phylogeography of alpine plants in the Sierra Nevada will be necessary before we understand all of the factors responsible for present distributions and predom- inant modes of speciation in the alpine flora of the range. Research Needs There is little doubt that the stability of the ecotone between alpine and treeline ecosystems in the Sierra Nevada and other high mountain regions has been and continues to be a function of complex interactions, with multiple drivers operating across diverse scales of time and space. This ecotone has been highly dynamic in the past and given the importance of temperature in controlling the elevation of treeline and higher alpine ecosystems, this ecotone and associated species are likely to be particularly sensitive to climate change in the future (Lloyd and Graum- lich 1997; Graumlich et al. 2005; Grabherr et al. 2010). Beyond treeline studies, the expansion of woody shrub species into alpine habitats has been shown to also be a sensitive indicator of potential climate change, with significant feedbacks on microclimate and soil ecosystems (Hallinger et al. 2010), as well as species facilitation (Callaway et al. 2002). 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L. STEPHENSON. 2000. Forest gradient response in Sierran landscapes: the physical template. Land- scape Ecology 15:603—620. VARGAS, P. 2003. Molecular evidence for multiple diversification patterns of alpine plants in Medi- terranean Europe. Taxon 52:463—476. WENT, F. W. 1948. Some parallels between desert and alpine floras in California. Madrono 9:241—249. . 1953. Annual plants at high altitudes in the Sierra Nevada, California. Madrono 12:109-114. WERNICKE, B., G. J. AXEN, AND J. K. SNOW. 1988. Basin and Range extensional tectonics at the latitude of Las Vegas, Nevada. Geological Society of America Bulletin 100:1738—1757. WOLFE, J. A., H. E. SCHORN, C. E. FOREST, AND P. MOLNAR. 1997. Paleobotanical evidence for high altitudes in Nevada during the Miocene. Science 276:1672-1675. 2011] RUNDEL: SIERRA NEVADA ALPINE FLORA 173 APPENDIX 1. Annotated checklist of the alpine flora of the Sierra Nevada, including all taxa reaching an elevation of 3500 m. Lower and upper elevations limits are those for all of California and taken from Baldwin et al. (2012). Growth form abbreviations are: P = erect broad-leaved perennial; G = geophytes; P-G = graminoid perennial; P-MAT = mat or cushion; A = annual; Q = aquatic perennial; SS = subshrub; S = woody shrub; and T = tree. Biogeographic relationships are abbreviated as follows: WIDE = widespread taxa present in many habitats or regions across North America and/or throughout the world; CORD = cordilleran taxa widespread in mountain regions of the western North America; S-C = Sierra/Cascade taxa with a Pacific Northwest distribution; INT = intermountain taxa present in the Great Basin; END = taxa endemic to the Sierra Nevada; and END-CAL = taxa endemic to California, as broadly defined in the text. Species names follow Baldwin et al. (2012). Specific or Lower Upper Growth Biogeographic Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship PTERIDOPHYTA Ophioglossaceae Botrychium crenulatum 1500 3600 P CORD Ophioglossaceae Botrychium lineare 2500 4000 P CORD Ophioglossaceae Botrychium paradoxum 4000 4200 P CORD Ophioglossaceae Botrychium simplex var. 1500 3800 Pp WIDE compositum Ophioglossaceae Botrychium tunux 3600 3600 led WIDE Pteridaceae Pellaea breweri 1500 3700 P INT Selaginellaceae Selaginella watsonii 1350 4100 P INT Woodsiaceae Athyrium distentifolium var. 1700 3700 P WIDE americanum Woodsiaceae Cystopteris fragilis 50 4100 i CORD Woodsiaceae Woodsia scopulina 1300 3500 P WIDE CONIFERAE Pinaceae Pinus albicaulis 2135 3700 T CORD Pinaceae Pinus balfouriana var. 2700 3700 i END austrina Pinaceae Pinus contorta subsp. 1525 3500 fi S-C murrayana Pinaceae Pinus flexilis 2600 3700 T CORD Pinaceae Tsuga mertensiana 1200 3500 T S-C MONOCOTY LEDONAE Alliaceae Allium obtusum var. obtusum 1500 3500 G INT Cyperaceae Carex albonigra 3000 4200 P-G CORD Cyperaceae Carex breweri 2000 3900 P-G S-C Cyperaceae Carex capitata 1200 3900 P-G WIDE Cyperaceae Carex congdonii 2600 3900 P-G END Cyperaceae Carex deflexa var. boottii 0 3800 P-G CORD Cyperaceae Carex douglasii 300 3800 P-G CORD Cyperaceae Carex filifolia var. erostrata 1500 3700 P-G CORD Cyperaceae Carex haydeniana 2400 4200 P-G CORD Cyperaceae Carex helleri 2400 4100 P-G S-C Cyperaceae Carex heteroneura 1300 4000 P-G INT Cyperaceae Carex hoodii 650 3600 P-G CORD Cyperaceae Carex incurviformis 3700 4000 P-G CORD Cyperaceae Carex jonesii 900 3500 P-G CORD Cyperaceae Carex lenticularis var. 0 3600 P-G CORD lipocarpa Cyperaceae Carex leporinella 1900 4000 P-G CORD Cyperaceae Carex mariposana 750 3600 P-G END-CAL Cyperaceae Carex multicostata 1900 3500 P-G CORD Cyperaceae Carex nigricans 1900 3700 P-G CORD Cyperaceae Carex phaeocephala 2500 4000 P-G CORD Cyperaceae Carex praeceptorium 2200 3500 P-G CORD Cyperaceae Carex proposita 3000 4100 P-G S-C Cyperaceae Carex rossii 0 3800 P-G CORD Cyperaceae Carex scirpoidea var. 2800 3700 P-G CORD pseudoscirpoidea Cyperaceae Carex specifica 1200 3500 P-G INT Cyperaceae Carex spectabilis 1800 3700 P-G CORD Cyperaceae Carex straminiformis 1700 4100 P-G S-C Cyperaceae Carex subfusca 700 3800 P-G INT Cyperaceae Carex subnigricans 2600 3800 P-G CORD Cyperaceae Carex tahoensis 3200 3700 P-G CORD 174 Higher level taxon Cyperaceae Cyperaceae Cyperaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaginaceae Liliaceae Melanthiaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae MADRONO APPENDIX |. _CONTINUED. Specific or Lower Upper infraspecific taxon elevation (m) elevation (m) Carex vernacula 1800 4000 Eleocharis quinqueflora 40 3600 Trichophorum clementis 2400 3600 Juncus bryoides 600 3600 Juncus drummondii 200 3500 Juncus mertensianus 1200 3500 Juncus mexicanus 0 3800 Juncus orthophyllus 1200 3500 Juncus parryi 2000 3800 Luzula divaricata 2100 3700 Luzula orestera 2700 3600 Luzula spicata 2900 3700 Luzula subcongesta 2000 3500 Triglochin palustris 2400 3500 Calochortus leichtlinii 1300 4000 Veratrum californicum var. 0 3500 californicum Agrostis idahoensis 0 3500 Agrostis pallens 200 3500 Agrostis scabra 100 3500 Agrostis thurberiana 1300 3500 Agrostis variabilis 1600 4000 Alopecurus aequalis var 50 3500 aequalis Bromus carinatus var. 0 3500 carinatus Bromus carinatus var. 0 3500 marginatus Bromus orcuttianus 560 3500 Bromus porteri 550 3500 Bromus richardsonii 1200 3600 Calamagrostis muiriana 2480 3900 Calamagrostis purpurascens 1300 4000 Deschampsia cespitosa 0 3820 subsp. cespitosa Elymus elymoides subsp. 295 4200 californicus Elymus multisetus 0 3800 Elymus scribneri 2900 4200 Elymus sierrae 1800 3530 Festuca brachyphylla subsp. 2800 4300 breviculmis Festuca minutiflora 2850 4050 Hordeum jubatum var. 20 3500 jubatum Koeleria macrantha 0 3840 Muhlenbergia richardsonis 1220 3670 Phleum alpinum 0 3700 Poa abbreviata subsp. 3300 3660 pattersonit Poa cusickii subsp. epilis 2400 3600 Poa cusickii subsp. 2100 3500 purpurascens Poa glauca subsp. rupicola 3300 4100 Poa keckii 3300 4340 Poa lettermanii 3500 4300 Poa secunda subsp. secunda 0 3900 Poa stebbinsii 2700 3700 Poa wheeleri 1300 3800 Stipa hymenoides 60 3500 Stipa kingii 2000 3650 Stipa nelsonii subsp. dorei 450 3500 Stipa occidentalis subsp. 1200 3500 pubescens Growth form P-G ror rg Rg tg ig bg hg by QAAAAAAAADADAOD an) ac lacldeclixcMaciacline| inc). \a-Sa°imia-|ta- MMM sc]ga°)iin-iin> MMMa-e>/ai tae) xc) og gela- ac) x-/bn-/ae) o7[la-]s Mela~ | lange ae) ec aclac exc > QQOQQQQQHQ0Q QAO AQAOQ AM AANAM ©Q AMAH0OO0H 0A 0 AHAMO0O0 [Vol. 58 Biogeographic relationship CORD WIDE END CORD CORD CORD WIDE CORD CORD S-C END WIDE S-C WIDE S-C CORD CORD CORD WIDE CORD CORD WIDE S-C S-C S-C CORD CORD END WIDE WIDE CORD CORD CORD INT WIDE CORD WIDE WIDE CORD WIDE CORD CORD S-C CORD END-CAL CORD CORD END CORD CORD END CORD CORD 2011] Higher level taxon Poaceae Poaceae Poaceae Poaceae Poaceae Themidaceae EUDICOTS Adoxaceae Apiaceae Apiaceae Apiaceae Apiaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae RUNDEL: SIERRA NEVADA ALPINE FLORA APPENDIX 1. Specific or infraspecific taxon Stipa pinetorum Stipa webberi Torreyochloa erecta Torreyochloa pallida var. pauciflora Trisetum spicatum Triteleia dudleyi Sambucus racemosa vat. melanocarpa Cymopterus cinerarius Oreonana clementis Podistera nevadensis Sphenosciadium capitellatum Achillea millefolium Ageratina occidentalis Agoseris aurantiaca var. aurantiaca Agoseris monticola Antennaria media Antennaria pulchella Antennaria rosea subsp. confinis Antennaria rosea subsp. rosea Antennaria umbrinella Arnica chamissonis Arnica lanceolata subsp. prima Arnica longifolia Arnica mollis Arnica ovata Artemisia arbuscula subsp. arbuscula Artemisia ludoviciana subsp. incompta Artemisia norvegica subsp. saxatilis Artemisia spiciformis Chaenactis alpigena Chaenactis douglasii var. douglasii Chrysothamnus viscidiflorus var. viscidiflorus Cirsium arizonicum var. arizonicum Cirsium occidentale var. venustum Cirsium scariosum var. americanum Crepis nana Ericameria bloomeri Ericameria discoidea Ericameria nauseosa var. Speciosa Ericameria parryi var. monocephala Ericameria suffruticosa Erigeron algidus Erigeron compositus Erigeron lonchophyllus Erigeron pygmaeus Erigeron vagus Lower elevation (m) 2000 1450 2000 0 1370 1200 1800 2100 1500 3000 CONTINUED. Upper elevation (m) 3900 3500 3500 3500 3900 3500 3600 3500 4000 4000 3500 3650 3700 3500 3800 3900 3700 3700 3700 3900 3500 3500 3500 3500 3600 3800 3500 3800 3700 3900 3500 4000 3500 3600 3500 4000 4000 3800 3500 3700 3800 3700 4300 3550 4100 4400 Growth form PPP @uaane eo 26 N wy Oy oC oy uv > > HH 175 Biogeographic relationship INT INT INT CORD WIDE END-CAL CORD INT END INT CORD WIDE CORD CORD S-C CORD INT WIDE CORD WIDE WIDE CORD CORD CORD CORD CORD INT WIDE CORD END-CAL CORD INT INT INT CORD CORD INT INT INT INT INT INT WIDE WIDE S-C INT 176 Higher level taxon Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae Brassicaceae APPENDIX |. Specific or infraspecific taxon Eriophyllum lanatum var. integrifolium Hazardia whitneyi var. whitneyi Hieracium nudicaule Hieracium triste Hulsea algida Hulsea vestita subsp. pygmaea Hymenoxys hoopesii Oreostemma alpigenum var. andersonii Oreostemma peirsonii Orochaenactis thysanocarpha Packera cana Packera werneriifolia Pyrrocoma apargioides Raillardella argentea Raillardella scaposa Senecio fremontii var. occidentalis Senecio integerrimus var. major Senecio pattersonensis Senecio scorzonella Senecio spartioides Solidago multiradiata Sphaeromeria cana Stenotis acaulis Tonestus peirsonii Cryptantha circumscissa vat. CIFCUMSCISSA Cryptantha circumscissa vat. rosulata Cryptantha glomeriflora Cryptantha humilis Cryptantha nubigena Hackelia micrantha Hackelia sharsmithii Nama densum Phacelia hastata subsp. compacta Phacelia mutabilis Phacelia ramosissima Anelsonia eurycarpa Boechera depauperata Boechera howellii Boechera inyoensis Boechera covillei Boechera inyoensis Boechera lemmonii Boechera lyallii Boechera paupercula Boechera repanda Boechera tiehmii Cardamine cordifolia Descurainia incana Draba albertina Draba breweri Draba cana Draba cruciata Draba densifolia Draba lemmonii MADRONO CONTINUED. Lower Upper elevation (m) elevation (m) 1400 3500 1200 3500 1800 3500 1650 3550 3000 4000 3200 3900 1500 3650 1200 3500 3000 3800 1600 3800 1200 3500 3000 3650 2200 3800 1800 3900 2000 3500 2800 4000 100 3600 3000 3700 1600 3500 1000 3500 1250 3950 1800 4000 1800 3600 2900 3700 150 3650 2950 3650 1800 3750 1700 3600 2400 3900 1200 3500 3150 3700 880 3560 1500 4000 900 3500 0 3800 1600 4100 3650 3900 1500 3800 1200 3500 2200 3500 1200 3500 2000 4350 2000 3900 2500 3700 1400 3600 3000 3600 600 3600 100 3500 900 3700 3100 4100 0 4100 2500 3963 1900 3650 3050 4000 Growth form iy eee Aan Dae lla > = - TUVUU DUE UU US UT >>Duv~TU> D> Sve AvA vy 4 a ti > v yuu > = [Vol. 58 Biogeographic relationship CORD END 2011] RUNDEL: SIERRA NEVADA ALPINE FLORA Li? APPENDIX |. CONTINUED. Specific or Lower Upper Growth Biogeographic Higher level taxon infraspecific taxon elevation (m) — elevation (m) form relationship Brassicaceae Draba lonchocarpa 2800 4000 P WIDE Brassicaceae Draba longisquamosa 3000 3900 P END Brassicaceae Draba novolympica 1500 3700 P-MAT CORD Brassicaceae Draba oligosperma 2000 3900 P-MAT CORD Brassicaceae Draba praealta 2500 4100 P WIDE Brassicaceae Draba sharsmithii 3300 3800 P END Brassicaceae Draba sierrae 3500 4114 P-MAT END Brassicaceae Draba subumbellata 3300 4100 P-MAT END-CAL Brassicaceae Erysimum capitatum var. 0 4000 P WIDE capitatum Brassicaceae Erysimum perenne 2000 4000 P S-C Brassicaceae Lepidium densiflorum 0 3500 Pp-A WIDE Brassicaceae Rorippa curvipes 100 3500 P-A CORD Brassicaceae Rorippa curvisiliqua 0 3500 A CORD Brassicaceae Streptanthus gracilis 2600 3600 A END Brassicaceae Streptanthus tortuosus 200 4100 P S-C Caryophyllaceae Cerastium beeringianum 2900 4300 P-MAT WIDE Caryophyllaceae Ereomogone kingii var. 2100 4050 P-MAT S-C glabrescens Caryophyllaceae Minuartia nuttallii var. 2600 3800 P-MAT S-C gracilis Caryophyllaceae Minuartia obtusiloba 3150 3700 P-MAT CORD Caryophyllaceae Minuartia rubella 2400 3800 P CORD Caryophyllaceae Minuartia stricta 3500 3900 Pp CORD Caryophyllaceae Sagina saginoides 1000 3800 P WIDE Caryophyllaceae Silene bernardina 1350 3600 P CORD Caryophyllaceae Silene sargentii 2400 3800 Pp S-C Caryophyllaceae Stellaria calycantha 1700 3800 Pp WIDE Chenopodiaceae Chenopodium atrovirens 300 3500 A CORD Chenopodiaceae Monolepis nuttalliana 0 3700 A CORD Crassulaceae Rhodiola integrifolia 1800 4000 Pr WIDE Crassulaceae Sedum obtusatum subsp. 1200 3700 P S-C obtusatum Ericaceae Cassiope mertensiana 1800 3505 S CORD Ericaceae Gaultheria humifusa 1350 4000 S CORD Ericaceae Kalmia polifolia subsp. 1000 3500 S CORD microphylla Ericaceae Phyllodoce breweri 1200 3500 S END-CAL Ericaceae Pterospora andromedea 60 3700 P WIDE Ericaceae Rhododendron columbianum 0 3630 S CORD Fabaceae Astragalus kentrophyta var. 2900 4000 P-MAT END-CAL danaus Fabaceae Astragalus kentrophyta var. 2700 3600 Er CORD tegetarius Fabaceae Astragalus lentiginosus var. 1250 3700 P INT ineptus Fabaceae Astragalus platytropus 2350 3500 P INT Fabaceae Astragalus purshii var. 1500 3650 P INT lectulus Fabaceae Astragalus whitneyi var. 1550 3500 P S-C whitneyi Fabaceae Lupinus adsurgens 1000 3500 P S-C Fabaceae Lupinus angustiflorus 1000 3500 P INT Fabaceae Lupinus argenteus var. 1500 3500 P INT meionanthus Fabaceae Lupinus argenteus var. 2500 3500 P INT montigenus Fabaceae Lupinus breweri var. breweri 1000 4000 P-MAT END-CAL Fabaceae Lupinus breweri var. 2500 4000 P-MAT END-CAL bryoides Fabaceae Lupinus breweri var. 2000 3500 P-MAT INT grandiflorus Fabaceae Lupinus covillei 2500 3500 P END Fabaceae Lupinus gracilentus 2500 3500 P END 178 MADRONO [Vol. 58 APPENDIX |. CONTINUED. Specific or Lower Upper Growth Biogeographic Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship Fabaceae Lupinus latifolius var. 1000 3500 iP S-C columbianus Fabaceae Lupinus latifolius var. 0 3500 ig END-CAL parishii Fabaceae Lupinus lepidus var. lobbii 2000 3500 P S-C Fabaceae Lupinus lepidus var. ramosus 3000 4000 P END-CAL Fabaceae Lupinus obtusilobus 2500 3500 P S-C Fabaceae Lupinus padre-crowleyi 2500 4000 P-MAT END-CAL Fabaceae Lupinus pratensis var. 1000 3500 P END-CAL pratensis Fabaceae Oxytropis borealis var. 3300 3900 P INT australis Fabaceae Oxytropis borealis var. 3300 3900 P CORD viscida Fabaceae Oxytropis parryi 3100 3800 P-MAT INT Fabaceae Trifolium kingii subsp. 2100 3500 P END dedeckerae Fabaceae Trifolium monanthum subsp. 1700 3900 P-MAT INT monanthum Gentianaceae Comastoma tenellum 3200 3900 A WIDE Gentianaceae Gentiana calycosa 1300 3900 P CORD Gentianaceae Gentiana newberryi var. 1500 4000 P. S-C tiogana Gentianaceae Gentianella amarella subsp. 1500 3500 A WIDE acuta Gentianaceae Gentianopsis holopetala 1800 4000 A S-C Grossulariaceae Ribes cereum var. inebrians 2100 3850 S INT Grossulariaceae Ribes montigenum 800 4000 S CORD Grossulariaceae Ribes velutinum 700 3500 S CORD Hydrangeaceae Jamesia americana 2070 3700 S INT Lamiaceae Monardella beneolens 2500 3600 SS END Lamiaceae Monardella linoides subsp. 1000 3500 SS INT sierrae Lamiaceae Monardella odoratissima 1000 3500 SS INT subsp. glauca Linaceae Linum lewisii 400 3657 P INT Montiaceae Calyptridium monospermum 300 3970 P INT Montiaceae Calyptridium roseum 1500 3800 A CORD Montiaceae Calyptridium umbellatum 240 4300 Pp CORD Montiaceae Claytonia nevadensis 2200 3500 P S-C Montiaceae Lewisia disepala 1300 3500 P END Montiaceae Lewisia glandulosa 3000 4000 P END-CAL Montiaceae Lewisia nevadensis 609 3596 P S-C Montiaceae Lewisia pygmaea 1700 4020 P CORD Montiaceae Lewisia triphylla 1300 3500 P CORD Montiaceae Montia chamissoi 1100 3700 P S-C Onagraceae Epilobium anagallidifolium 1500 4500 Pp WIDE Onagraceae Epilobium ciliatum subsp. 0 4000 P CORD ciliatum Onagraceae Epilobium ciliatum subsp. 0 3500 P WIDE glandulosum Onagraceae Epilobium clavatum 1200 4200 P CORD Onagraceae Epilobium glaberrimum 1200 3800 P CORD subsp. fastigiatum Onagraceae Epilobium hallianum 100 3700 P CORD Onagraceae Epilobium hornemannii 1200 3900 P WIDE subsp. hornemannii Onagraceae Epilobium obcordatum 1700 4000 P S-C Onagraceae Epilobium oregonense 1200 3500 P CORD Onagraceae Epilobium saximontanum 1400 3500 P CORD Onagraceae Gayophytum decipiens 1800 4200 A INT Onagraceae Gayophytum diffusum subsp. 800 3700 A S-C diffusum 2011] Higher level taxon Onagraceae Onagraceae Onagraceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Orobanchaceae Parnassiaceae Phrymaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Plantaginaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polemoniaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae RUNDEL: SIERRA NEVADA ALPINE FLORA APPENDIX 1. CONTINUED. Specific or Lower Upper infraspecific taxon elevation (m) elevation (m) Gayophytum diffusum subsp. 800 3700 parviflorum Gayophytum racemosum 1000 4000 Gayophytum ramosissimum 500 3500 Castilleja applegatei subsp. 1900 3600 pallida Castilleja applegatei subsp. 300 3600 pinetorum Castilleja lemmonii 1550 3700 Castilleja miniata subsp. 1500 3500 miniata Castilleja nana 2400 4200 Pedicularis attollens 1200 4000 Pedicularis groenlandica 1000 3600 Pedicularis semibarbata 1500 3500 Parnassia palustris 0 3600 Mimulus suksdorfii 1100 4000 Callitriche palustris 0 4000 Collinsia parviflora 800 3500 Collinsia torreyi var. wrightii 800 4000 Penstemon davidsonii 2000 3750 Penstemon heterodoxus var. 2700 3900 heterodoxus Penstemon newberryi var. 1000 3700 newberryi Penstemon procerus var. 2100 3600 formosus Penstemon roezlii 300 3500 Penstemon rostriflorus 500 3500 Penstemon rydbergii var. 1000 3600 oreocharis Penstemon speciosus 850 3800 Veronica wormskjoldii 1500 3500 Collomia linearis 600 3650 Gymnosteris parvula 2400 3700 Ipomopsis congesta subsp. 1500 3700 montana Leptosiphon oblanceolatus 2800 3700 Linanthus pungens 1700 4000 Phlox condensata 2000 4000 Phlox diffusa 1100 3600 Phlox dispersa 3600 4200 Phlox pulvinata 3300 4300 Polemonium eximium 3000 4200 Polemonium pulcherrimum 2400 3700 var. pulcherrimum Eriogonum gracilipes 2900 3900 Eriogonum incanum 2100 4000 Eriogonum lobbii 1600 3800 Eriogonum nudum var. 2800 3800 scapigerum Eriogonum ovalifolium var. 3000 3600 caelestinum Eriogonum ovalifolium var. 1700 4200 nivale Eriogonum polypodum 2800 3500 Eriogonum rosense var. 2300 4000 rosense Eriogonum spergulinum var. 1300 3500 pratense Eriogonum umbellatum var. 3000 3600 covillei Eriogonum wrightii var. 3500 3600 olanchense Growth form A tT > > >PrOruwu yw ys UT CU P-MAT P-MAT P-MAT P-MAT ~ Su> Uppy VAL - 179 Biogeographic relationship INT CORD INT INT INT S-C INT END-CAL S-C S-C S-C WIDE INT WIDE WIDE INT INT S-C INT INT INT INT INT INT WIDE WIDE CORD S-C END CORD CORD CORD END CORD END S-C END-CAL INT S-C END END INT END INT END END-CAL END 180 MADRONO [Vol. 58 APPENDIX |. CONTINUED. Specific or Lower Upper Growth Biogeographic Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship Polygonaceae Oxyria digyna 1800 4000 P WIDE Polygonaceae Rumex californicus 0 3500 P CORD Polygonaceae Rumex paucifolius 1500 4000 P S-C Polygonaceae Rumex salicifolius 0 3500 P INT Polygonaceae Rumex utahensis 1000 3500 P CORD Primulaceae Androsace septentrionalis 2700 3600 P-A WIDE Primulaceae Dodecatheon redolens 2400 3600 P INT Primulaceae Dodecatheon subalpinum 2100 4000 P END Primulaceae Primula suffrutescens 2000 4200 P END-CAL Pteridaceae Pellaea breweri 1500 3700 P INT Ranunculaceae Aconitum columbianum 300 3500 | CORD subsp. columbianum Ranunculaceae Aquilegia pubescens 2600 3650 P END Ranunculaceae Delphinium polycladon 2200 3600 P END-CAL Ranunculaceae Ranunculus alismifolius var. 1400 3600 P S-C alismellus Ranunculaceae Ranunculus eschscholtzii var. 2200 3600 P S-C eschscholtzii Ranunculaceae Ranunculus eschscholtzii var. 2700 4300 P END-CAL oxynotus Ranunculaceae Ranunculus glaberrimus 1200 3600 P CORD Ranunculaceae Thalictrum alpinum 2900 3700 P WIDE Ranunculaceae Thalictrum sparsiflorum 1400 3500 P CORD Rosaceae Dasiphora fruticosa 2000 3600 S WIDE Rosaceae Drymocaulis lactea var. 1800 3700 P INT lactea Rosaceae Drymocaulis pseudorupestris 3200 3900 P CORD var. crumiana Rosaceae Drymocaulis pseudorupestris 2300 3500 leg CORD var. saxicola Rosaceae Holodiscus discolor var. 1159 4000 S CORD microphyllus Rosaceae Ivesia gordonii var. 1800 3500 P INT ursinorum Rosaceae Ivesia lycopodioides subsp. 3000 4000 P END-CAL lycopodioides Rosaceae Ivesia lycopodioides subsp. 3000 4115 P END-CAL scandularis Rosaceae Ivesia muirii 2900 4000 P END Rosaceae Ivesia pygmaea 2700 4000 P END Rosaceae Ivesia santolinoides 1500 3600 P END-CAL Rosaceae Ivesia shockleyi 2700 4000 P-MAT INT Rosaceae Potentilla breweri 1500 3700 P S-C Rosaceae Potentilla flabellifolia 1700 3700 P S-C Rosaceae Potentilla bruceae 1200 3700 P INT Rosaceae Potentilla glaucophylla var. 2600 3500 P WIDE glaucopylla Rosaceae Potentilla gracilis var. 800 3500 P INT fastigiata Rosaceae Potentilla jepsonii 2700 3800 P INT Rosaceae Potentilla pensylvanica 2700 3800 P WIDE Rosaceae Potentilla pseudosericea 3200 4300 P END-CAL Rosaceae Potentilla wheeleri 1800 3500 P END-CAL Rosaceae Sibbaldia procumbens 1820 3700 | a WIDE Rosaceae Sorbus californica 1200 4300 S INT Rubiaceae Galium bifolium 1500 3700 A CORD Rubiaceae Galium grayanum var. 1830 3500 P S-C grayanum Rubiaceae Galium hypotrichium subsp. 3000 4200 P END-CAL hypotrichium Rubiaceae Galium hypotrichium subsp. 2650 3880 P END subalpinum Salicaceae Salix brachycarpa var. 3200 3500 S CORD brachycarpa 2011] RUNDEL: SIERRA NEVADA ALPINE FLORA 18] APPENDIX |. CONTINUED. Specific or Lower Upper Growth Biogeographic Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship Salicaceae Salix eastwoodiae 1600 3800 S CORD Salicaceae Salix geyeriana 1450 3600 S CORD Salicaceae Salix lemmonii 1400 3500 S CORD Salicaceae Salix nivalis 3100 3500 S CORD Salicaceae Salix orestera 1100 4000 S S-C Salicaceae Salix petrophila 1670 4000 S CORD Salicaceae Salix planifolia 2500 4000 S WIDE Saxifragaceae Heuchera rubescens 1000 4000 P CORD Saxifragaceae Lithophragma glabrum 0 3750 P CORD Saxifragaceae Micranthes aprica 1600 3600 P CORD Saxifragaceae Micranthes bryophora 1600 3500 P CORD Saxifragaceae Micranthes nidifica 1000 3500 lg CORD Saxifragaceae Micranthes tolmiei 1980 3596 eg CORD Saxifragaceae Pectiantia breweri 1500 3500 P S-C Saxifragaceae Saxifraga hyperborea 3000 4500 P WIDE Valerianaceae Valeriana californica 1500 3700 P INT Violaceae Viola adunca 0 3570 P WIDE Violaceae Viola bakeri 900 3800 P INT Violaceae Viola macloskeyi 609 3600 P WIDE Violaceae Viola pinetorum subsp. 1981 3700 P END-CAL grisea Violaceae Viola purpurea subsp. 1400 3598 le END-CAL mesophyta APPENDIX 2. Annotated checklist of the flora of the Sierra Nevada with an upper elevational limit of 3300— 3499 m. Abbreviations as in Appendix 1. Species names follow Baldwin et al. (2012). Higher level Specific or Lower Upper Growth Biogeographic taxon infraspecific taxon elevation (m) elevation (m) form relationship PTERIDOPHYTA Aspleniaceae Asplenium septentrionale 2500 3350 P WIDE Ophioglossaceae Botrychium lunaria 2300 3400 Pp WIDE Pteridaceae Adiantum aleuticum 0 3400 | WIDE Pteridaceae Aspidotis densa 100 3400 Pp CORD Pteridaceae Cryptogramma acrostichoides 1400 3400 P INT CONIFERAE Pinaceae Pinus monticola 150 3400 T S-C MONOCOTS Alliaceae Allium validum 1200 3400 G CORD Cyperaceae Carex abrupta 1200 3450 P-G S-C Cyperaceae Carex aurea 1100 3300 P-G WIDE Cyperaceae Carex buxbaumii 0 3300 P-G WIDE Cyperaceae Carex davyi 1400 3300 P-G S-C Cyperaceae Carex disperma 1100 3400 P-G WIDE Cyperaceae Carex fissuricola 1500 3300 P-G CORD Cyperaceae Carex fracta 250 3300 P-G S-C Cyperaceae Carex illota 2100 3400 P-G CORD Cyperaceae Carex integra 800 3400 P-G S-C Cyperaceae Carex microptera 1500 3400 P-G CORD Cyperaceae Carex pellita 60 3300 P-G WIDE Cyperaceae Carex petasata 600 3400 P-G CORD Cyperaceae Carex preslii 1800 3400 P-G CORD Cyperaceae Carex simulata 0 3300 P-G CORD Cyperaceae Carex tiogana 3100 3350 P-G S-C Cyperaceae Carex utriculata 0 3400 P-G WIDE Cyperaceae Carex vesicaria 0 3300 P-G WIDE Cyperaceae Carex whitneyi 1200 3400 P-G S-C 182 MADRONO [Vol. 58 APPENDIX 2. CONTINUED. Higher level Specific or Lower Upper Growth Biogeographic taxon infraspecific taxon elevation (m) elevation (m) form relationship Cyperaceae Eleocharis acicularis var. acicularis 0 3300 P-G WIDE Cyperaceae Eleocharis acicularis var. gracilescens 0 3300 P-G WIDE Cyperaceae Eleocharis suksdorfiana 0 3400 P-G CORD Cyperaceae Eriophorum criniger 2000 3350 P-G S-C Iridaceae Tris missouriensis 900 3400 G CORD Juncaceae Juncus hemiendytus var. abjectus 1400 3400 P-G INT Juncaceae Juncus nevadensis subsp. nevadensis 1200 3300 P-G CORD Juncaceae Luzula parviflora var. parviflora 1000 3300 P-G WIDE Liliaceae Lilium kelleyanum 2200 3300 G END Orchidaceae Platanthera dilatata var. leucostachys 0 3400 G CORD Orchidaceae Platanthera sparsiflora 100 3400 G CORD Orchidaceae Spiranthes romanzoffiana 0 3300 P WIDE Poaceae Agrostis humilis 1500 3350 P-G CORD Poaceae Bromus suksdorfii 1250 3300 P-G S-C Poaceae Calamagrostis canadensis var. 1500 3400 P-G WIDE canadensis Poaceae Calamagrostis canadensis vat. 1500 3400 P-G WIDE langsdorfii Poaceae Calamagrostis stricta subsp. inexpansa 0 3400 P-G WIDE Poaceae Calamagrostis stricta subsp. stricta 1500 3350 P-G WIDE Poaceae Danthonia intermedia var. intermedia 1460 3450 P-G WIDE Poaceae Elymus trachycaulus subsp. 0 3400 P-G WIDE trachycaulus Poaceae Hordeum brachyantherum subsp. 0 3400 P-G CORD brachyantherum Poaceae Hordeum brachyantherum subsp. 0 3400 P-G END-CAL californicum Poaceae Melica bulbosa 0 3400 P-G CORD Poaceae Melica stricta 1200 3350 P-G INT Poaceae Muhlenbergia filiformis 150 3350 A CORD Poaceae Muhlenbergia montana 1640 3420 P-G WIDE Poaceae Stipa nevadensis 1000 3450 P-G CORD Poaceae Stipa occidentalis subsp. californica 150 3450 P-G CORD Poaceae Stipa occidentalis subsp. occidentalis 1200 3450 P-G CORD Poaceae Trisetum wolfii 1740 3300 P-G CORD EUDICOTS Apiaceae Angelica lineariloba 1700 3300 P END-CAL Apiaceae Ligusticum grayi 1000 3300 P CORD Apiaceae Lomatium torreyi 1100 3300 P END Apiaceae Perideridia parishii subsp. latifolia 2000 3400 P S-C Asteraceae Agoseris parviflora 1400 3400 P CORD Asteraceae Artemisia cana subsp. bolanderi 1200 3300 S S-C Asteraceae Artemisia dracunculus 0 3400 ig INT Asteraceae Chaenactis douglasii var. alpina 3000 3400 P-MAT CORD Asteraceae Crepis acuminata 1000 3300 P CORD Asteraceae Crepis intermedia 800 3300 CORD Asteraceae Ericameria parryi var. aspera 1900 3300 SS INT Asteraceae Erigeron barbellulatus 2100 3300 P END-CAL Asteraceae Erigeron clokeyi var. pinzliae 2200 3400 P INT Asteraceae Erigeron coulteri 1900 3400 P CORD Asteraceae Erigeron elmeri 1300 3300 P END Asteraceae Erigeron glacialis var. glacialis 1300 3400 E CORD Asteraceae Erigeron tener 2300 3400 P-MAT CORD Asteraceae Helenium bigelovii 0 3400 P S-C Asteraceae Hieracium albiflorum 0 3300 P WIDE Asteraceae Hieracium horridum 1350 3300 P S-C Asteraceae Hulsea vestita subsp. vestita 2400 3350 P END Asteraceae Microseris nutans 1000 3400 P CORD Asteraceae Nothocalais alpestris 1300 3400 P S-C Asteraceae Packera pauciflora 1800 3300 P WIDE Asteraceae Senecio triangularis 100 3300 P CORD Asteraceae Tetradymia canescens 1000 3400 S CORD Asteraceae Tonestus eximius 1800 3300 P CORD Asteraceae Wyethia mollis 900 3400 P S-C 2011) RUNDEL: SIERRA NEVADA ALPINE FLORA 183 APPENDIX 2. CONTINUED. Higher level Specific or Lower Upper Growth Biogeographic taxon infraspecific taxon elevation (m) elevation (m) form relationship Boraginaceae Cryptanrtha watsonii 1250 3300 A CORD Boraginaceae Cryptantha confertiflora 1050 3350 P INT Boraginaceae Lappula redowskii 1300 3300 A WIDE Boraginaceae Mertensia ciliata 1300 3380 P S-C Boraginaceae Phacelia bicolor 700 3400 A INT Boraginaceae Phacelia eisenii 1300 3400 A END Boraginaceae Phacelia orogenes 2060 3400 A END Boraginaceae Plagiobothrys hispidulus 1200 3400 A CORD Boraginaceae Plagiobothrys torreyi var. diffusus 1200 3400 A END-CAL Brassicaceae Barbarea orthoceras 0 3400 B/P WIDE Brassicaceae Boechera calderi 2050 3350 P CORD Brassicaceae Boechera davidsonii 1200 3400 PB S-C Brassicaceae Boechera pygmaea 2600 3400 P END Brassicaceae Boechera stricta 1800 3400 P CORD Brassicaceae Cardamine oligosperma 50 3300 A/B CORD Brassicaceae Descurainia californica 1700 3400 A/B CORD Brassicaceae Draba asterophora 2600 3300 P INT Brassicaceae Physaria occidentalis 600 3350 PE INT Caprifoliaceae Lonicera conjugialis 140 3300 S S-C Caprifoliaceae Symphicarpos rotundifolius var. parishii 1100 3300 S INT Caryophyllaceae Ereomogone congesta var. 1200 3300 P INT subfrutescens Ericaceae Arctostaphylos patula 750 3350 S CORD Ericaceae Arctostaphylos uva-ursi 2400 3300 S WIDE Ericaceae Vaccinium caespitosum 0 3400 S WIDE Ericaceae Vaccinium uliginosum subsp. 0 3400 S WIDE occidentale Fabaceae Astragalus bolanderi 1400 3300 P INT Fabaceae Astragalus raventii 3400 3450 P END Fabaceae Trifolium monanthum subsp. tenerum 1600 3300 P END Fagaceae Chrysolepis sempervirens 700 3300 S S-C Gentianaceae Frasera puberulenta 1700 3400 P END-CAL Gentianaceae Gentianopsis simplex 1200 3400 P INT Grossulariaceae Ribes inerme var. inerme 1200 3300 S CORD Lamiaceae Monardella breweri subsp. lanceolata 0 3400 A INT Montiaceae Claytonia megarhiza 2600 3300 P CORD Montiaceae Lewisia leana 1300 3350 P S-C Onagraceae Chamerion angustifolium subsp. 0 3300 P WIDE circumvagum Orobanchaceae Castilleja arachnoidea 1300 3300 P INT Orobanchaceae Castilleja linariifolia 1000 3350 P INT Orobanchaceae Castilleja peirsonii 1500 3400 P S-C Orobanchaceae Castilleja pilosa 1200 3400 P INT Orobanchaceae Castilleja praeterita 2200 3400 ley END Orobanchaceae Orobanche fasciculata 0 3300 P WIDE Papaveraceae Dicentra uniflora 1000 3300 Pp. CORD Phrymaceae Mimulus breweri 1200 3400 A CORD Phrymaceae Mimulus nanus var. mephiticus 1520 3445 A S-C Phrymaceae Mimulus tilingii 1400 3400 P CORD Plantaginaceae Penstemon caesius 1800 3400 SS END-CAL Plantaginaceae Veronica americana 0 3300 P WIDE Polemoniaceae Ipomopsis aggregata subsp. bridgesii 1800 3300 P END Polemoniaceae Microsteris gracilis 0 3300 A WIDE Polemoniaceae Navarretia breweri 1000 3300 A CORD Polemoniaceae Polemonium occidentale subsp. 900 3300 P WIDE occidentale Polygonaceae Eriogonum latens 2600 3400 P END-CAL Polygonaceae Eriogonum microthecum var. alpinum 2500 3300 SS END-CAL Polygonaceae Eriogonum microthecum var. ambiguum 1100 3300 SS INT Polygonaceae Eriogonum saxatile 800 3400 P-MAT INT Polygonaceae Eriogonum spergulinum vat. 1300 3400 A INT reddingianum Polygonaceae Eriogonum wrightii var. subscaposum 200 3400 P-MAT INT 184 Higher level taxon Polygonaceae Polygonaceae Potamogetonaceae Primulaceae Ranunculaceae Ranunculaceae Ranunculaceae Ranunculaceae Rhamnaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Salicaceae Salicaceae Scrophulariaceae MADRONO APPENDIX 2. Specific or infraspecific taxon Polygonum polygaloides subsp. kelloggii Polygonum shastense Potamogeton robbinsii Dodecatheon alpinum Anemone drummondii Aquilegia formosa Caltha leptosepala var. biflora Delphinium nuttallianum Ceanothus cordulatus Amelanchier utahensis Fragaria virginiana Geum macrophyllum var. perincisum Horkelia fusca subsp. parviflora Ivesia gordonii var. alpicola Ivesia saxosa Rosa woodsii subsp. gratissima Spiraea splendens Salix jepsonii Salix scouleriana Limosella acaulis CONTINUED. Lower Upper elevation (m) elevation (m) 1500 3300 2100 3400 1600 3300 1700 3400 1200 3350 0 3300 900 3300 300 3300 365 3365 200 3400 1200 3300 1000 3300 1400 3300 2100 3300 900 3300 800 3400 548 3400 1000 3400 l 3400 ) 3300 [Vol. 58 Growth Biogeographic form P NNnNNANDODIadDAWMN DI VIdIVOV a Q relationship CORD S-C WIDE CORD S-C S-C CORD CORD INT INT WIDE WIDE CORD INT INT INT S-C S-C CORD WIDE MADRONO, Vol. 58, No. 3, pp. 185-189, 2011 A NEW SPECIES OF ASTRAGALUS (FABACEAE) FROM THE WASATCH MOUNTAINS OF UTAH BETH LOWE CORBIN Owyhee Field Office, Bureau of Land Management, 20 First Ave. W, Marsing, ID 83639 ecorbin@blm.gov ABSTRACT Astragalus kelseyae B.L. Corbin, sp. nov. is described as a new species from the central Wasatch Mountains, where it is known from only one small occurrence in Weber County, Utah. It grows in shale talus within the Gambel oak and bigtooth maple shrubland. Key Words: Astragalus, Fabaceae, rare plant, Utah, Wasatch Mountains. A new species of Astragalus is described from a single population in the foothills of the Wasatch Mountains in Weber County, Utah. This distinc- tive milkvetch’s fruit shape and texture resemble some forms of A. Jentiginosus Douglas ex Hooker, but plants differ in having a branching, subterranean caudex, a smaller fruit beak, and generally larger (and fewer) flowers and fruit. Its humistrate growth form and fruit shape resemble A. amblytropis Barneby, but it has firmer fruits and larger flowers. Its large pods are similar to A. megacarpus (Nutt.) A. Gray, and flowers and leaflets are similar to A. beckwithii Torr. & A. Gray, but it differs from both species by having bilocular fruit, shorter leaves, and larger flowers. It differs significantly from each of those species by having dolabriform (malpighian) hairs. TAXONOMIC TREATMENT Astragalus kelseyae B.L. Corbin, sp. nov. (Fig. 1) — Type: USA, Utah, Weber Co., talus slope above Ogden, TON, RIW, Sec 26, SLM, 41°14'N, 111°55'W, 1625 m (5340 ft) elevation, 28 May 2010, Beth Lowe Corbin 1292 (holo- type: UT; isotypes: NY, CIC, UVSC, to be distributed). Planta similis A. /entiginosus Douglas ex Hooker var. negundo S.L. Welsh & N.D. Atwood et A. amblytropis Barneby in legumina biloculares et e var. negundo in caudices subteranneis elongatis et e ambo in legumina cartilagineis nec chartaceis et pubescentiis dolabriformis differt. Perennial herb from branched, subterranean, woody caudex branches. Above-ground stems 10-20 cm long, prostrate (Fig. 2). Leaves humi- strate, 3.05.2 cm long, 1.5—3.0 cm wide, with (5) 7-11 (13) leaflets, the terminal jointed. Leaflets widely elliptical, 8-15 mm long, 6-12 mm wide, tips rounded to obtuse, more or less alternate on the rachis. Leaves and stems silvery green with appressed dolabriform hairs about 0.5—0.8 mm long. Stipules free, triangular, 4-5 mm _ long. Inflorescence 2—7 flowered, congested, not much elongating in fruit. Peduncle 13—25 mm; flowering axis 5-10 mm. Calyx 13—16 mm long, 3.5—4.5 mm wide, with narrow teeth 3-4 mm long; calyx pinkish, with light and dark hairs. Petals white, with keel tip slightly purple. Banner 22—26 mm long, 9-11 mm wide, bent midway at about a 120° angle. Wings 19-21 mm long, narrow, slightly cupped. Keel 18-19 mm long. Fruit firm, cartilaginous, fleshy, inflated, bilocular, humi- strate, sessile, with narrow and shallow dorsal and ventral grooves, not curved, sometimes red mottled. Fresh pod 35-53 mm long, 18—30 mm thick, 10-17 mm tall (dorsiventrally compressed), with a small beak 3-6 mm long, and _ short, dolabriform hairs. Dehiscence through the beak, after separation. Paratypes (topotypes): 19 May 2009 Beth Lowe Corbin 1235 (BRY), 4 September 2011 Beth Lowe Corbin 1523 (UTC — to be distributed). DISTRIBUTION AND HABITAT Astragalus kelseyae grows on the lower, west/ southwest-facing slope of the central Wasatch Mountains, on talus openings within Quercus gambelii Nutt. (Gambel oak) and Acer grand- identatum Nutt. (bigtooth maple) shrublands. The talus consists of fine-textured Ophir shale on about 50-60% slope, at about 1625 m elevation. This habitat is just above the old shoreline of the Pleistocene Lake Bonneville. Precipitation is about 51—64 cm (20-25 inches) per year. The site 1s within the Uinta-Wasatch- Cache National Forest. Astragalus kelseyae 1s known only from the type locality, where about 150-200 plants were seen in a localized area of about 0.1 ha. It was first found in 2009, and revisited in 2010 and 2011. A popular. hiking trail bisects the occurrence, and additional undesig- nated trails occur within the habitat. The geology of the Wasatch Mountains is a complicated mix of sedimentary, metamorphic (such as quartzite), and igneous deposits (Yonkee and Lowe 2004). Ophir shale is a Paleozoic era 186 MADRONO [Vol. 58 Fic. 1. sedimentary deposit. Bands of Ophir shale and other similar talus types occur in scattered locations across the Wasatch Front, and addi- tional occurrences of Astragalus kelseyae may be found in the future. The talus microsite is very open, with low cover by associates, including Scutellaria angustifolia Astragalus kelseyae habit, flower detail, pod cross section, and leaf hair detail. Pursh subsp. micrantha Olmstead, Asclepias asperula (Decne.) Woodson, Apocynum androsae- — mifolium L., Epilobium canum (Greene) P.H. | Raven subsp. garrettii (A. Nelson) P.H. Raven, | Hedysarum boreale Nutt., Erysimum capitatum | (Douglas) Greene, Eriogonum umbellatum Torr., | Pseudoroegneria spicata (Pursh) A. Love, Ame- | 2011] CORBIN: A NEW ASTRAGALUS FROM UTAH 187 FIG. 2. Astragalus kelseyae in its shale talus habitat. lanchier utahensis Koehne, and Phacelia hastata Douglas ex Lehm. Although no weeds occur directly with the Astragalus, several weedy species occur in the vicinity, including J/satis tinctoria L., Linaria dalmatica (L.) Mill., Euphorbia myrsinites L., and Bromus tectorum L., and pose a threat to this species. RELATIONSHIPS Astragalus kelseyae appears to have similarities to A. lentiginosus, A. amblytropis, A. megacarpus, and A. beckwithii, but differs significantly from each (S. Welsh, Brigham Young Univ., personal communication). Its pod resembles A. lentigino- sus var. negundo with a large, bilocular fruit, but differs in having a less prominent beak, generally wider fruit, fewer flowers, a branched, subterra- nean caudex, and dolabriform hairs. It is similar to A. amblytropis in having a subterranean caudex, humistrate stems, leaves, and fruit, and somewhat similar fruit shape, but differs in having firmer pods, larger flowers, free stipules, -and dolabriform hairs. It superficially appears similar to A. megacarpus or A. oophorus S. _Watson with its large pods, and A. beckwithii -with its white flowers, but differs from these species in the section Megacarpi by both its bilocular fruit and dolabriform hairs. Thus, A. kelseyae differs from each of these species by the presence of dolabriform hairs, and other charac- teristics as shown in Table 1. The new species’ resemblance to A. /entiginosus is likely due to independent parallel evolution, rather than a close relationship; the evolution of a fruit septum (and dolabriform hairs) does not necessarily imply shared ancestry with other species with these characters (J. Alexander, Utah Valley Univ., personal communication). Dolabriform hairs have apparently arisen independently in several sections within genus, but none of the other species with dolabriform hairs have large, bilocular pods, relatively few, large flowers, and prostrate stems. The combination of characteris- tics present in A. ke/sevyae appears unique. Astragalus lentiginosus, with its plethora of varieties, 1s widespread, including Weber Co., Utah; var. negundo is known from Box Elder Co. (which is adjacent to the north side of Weber Co.) and Millard and Tooele counties (southwest of Weber Co.), Utah, so its range somewhat overlaps that of A. kelseyae (Welsh 2007). Astragalus amblytropis is limited to the vicinity of Challis, in Custer and Lemhi counties, Idaho, some 290 miles north of A. ke/seyvae (Welsh 2007). Astragalus megacarpus 1s known from Wyoming MORPHOLOGICAL COMPARISON OF ASTRAGALUS KELSEYAE AND SIMILAR SPECIES. Measurements (except A. kelseyae) come from Barneby 1989, Welsh on TABLE 1. 88 MADRONO [Vol. 58 (at least 50 miles east of this occurrence), central 26 od 6 ae Uo) of) oO : Go ee eee rey: sone Utah (about 120 miles south), Nevada, and Gos Sf 8S 8s Oc ; eee: as 2 O ee 5 Colorado, while A. beckwithii occurs in Weber a 0° EAE Aas 3 = : = Z = 0 © 5 Seiten fi aise Co. and generally to the west in Utah, and in fae) : Ic a 5 ee ee eo Idaho, Nevada, Oregon, Washington, and British S| 2S8i 582 B57 FEL] Columbia (Albee et al. 1988; Welsh et al. 2008; — oe > iol Swe OW aI(PEOEREES o8y Sure Shultz et al. 2010; USDA 2010). Astragalus Gs owns — = . Seleaceec o fF Tess, = oophorus occurs mostly in the southern Great — qc ba — > ° . 5 se Pore ee. Wee, geet ae Basin, east into western Colorado, but is also . far} mo oe i? wa = recorded from northwest Utah (Box Elder and fan} on . ; Tooele counties) (Welsh et al. 2008; USDA 2010). Bs 9 ae a ae Soe Fes & Thus, the location of A. kelseyae is at the east edge = 9 ae +6 ON rs wn ey Oo7r& KEY TO SIMILAR SPECIES cee: ome Zo S86 pee (adapted from Barneby 1989 and Welsh 2007) On = OQ, mM WH 1. Hairs dolabriform; pods fully bilocular ...A. kelseyae o 8 = fon) é : P : : i oh. - — 1’. Hairs basifixed; pods unilocular or bilocular 7 Sea 6... Be oA 5 ns 2. Pods unilocular ™ > ~ sce S 5 c os c S So 2 bby 3. Pod 1.5—3.7 cm long, valves leathery. . . = BS %Rh560 FS pee eae ye ee eee eee A. beckwithii | S = 5 « a..8 & 5 a = 3’. Pod (2) 3-6 (7.5) cm long, valves papery = v . A oS Ss wee. = Oe 4. Plants subacaulescent, stems 1-— i ees na Bas Z eG of b eC TONS yiext cen eee A, megacarpus meas 8 7 = 38 a ce SE S 4’. Plants caulescent, stems mostly 5— @ : o ee oe oe at 20 cmon? ta.e foe ee A. oophorus ere “a a = 2’. Pods bilocular eo = Z . 4 . Sp - a a 5. Stipules connate; caudex deeply sub- : Ces : oe ae . = =z 6yaea E LORRAINE Ato sre ane ee A. amblytropis > a OO Aa & bh : : i f2 ¢ a s & = a. 5’. Stipules free; caudex superficial (near SS Oo aie SB aT £2 SOil Surlace): = 4..°7 eee ee A. lentiginosus aS AN a : ms 2 ia 2 do 5 ns nm Pos Ot N 0) = op 80 ne | a | “oO Q = 7 5° oa = on On ase oO. a Res ACK Se a ;S a0, Ff oO Bo CKNOWLEDGMENTS aS Dn Ymadiad K Oh fice ee ; : ; A 5 fs S B ears a2 St mI = I thank Stanley Welsh (Brigham Young University, = @ Go = FREL SOO mo BRY) for detailed information, the Latin diagnosis, and O79 a 5 mn 2 access to the herbarium. I also thank Stephen Clark (Weber State University, WSCO), Don Mansfield Prt y . . a ~< (College of Idaho, CIC), Michael Piep (Utah State S ai i= re NaTi-=| University, UTC) and Ann Kelsey (University of Utah, = | qaq59S FS 3 oak . ew) SxS, 2eB = = ty ip SO ob UT) for access to collections. Jason Alexander (Utah ~_ 2 ot fw ao o & < Soh is a S Valley University) provided helpful comments and S c 5 = ; HAE 5 2 o = oo ffi 7 insights. Dan Scott produced the illustration, which was s NJBPU Sm we gunk. 6 5- funded by the Intermountain Region of the U.S. Forest = S2H2 sees aud adaat 5 ear = SERB F RSA SBSNS EL KMS Service, thanks to Teresa Prendusi, who also reviewed a co) = G c - s : 7 = Coe c Z Sayer draft of this article, as did Don Mansfield. This paper was _| ao) also improved by reviews from Stanley Welsh and Leila | = Shultz; however, any mistakes are all mine. lan} (cD) > 3 oc} 18 = S| 88 : 3 al | 9 5 on LITERATURE CITED ia) Nn 0) =— @ = S| 86 = 2 ogee = =|3 0 S 2 = 3 ALBEE, B. J., L. M. SHULTZ, AND S. GOODRICH. 1988. Atlas D UIAn [Ly [Ly = 4 of the vascular plants of Utah. Utah Museum of | 2011] Natural History, University of Utah, Salt Lake City, Li BARNEBY, R. C. 1989. Fabaceae. Pp. 12-267 in A. Cronquist, A. H. Holmgren, N. H. Holmgren, J. L. Reveal, and P. K. Holmgren (eds.), Intermountain flora, Vol. 3, pt. B: Fabales. The New York Botanical Garden, Bronx, NY. SHULTZ, L. M., R. D. RAMSEY, W. LINDQUIST, AND C. GARRARD. 2010. Digital atlas of the vascular plants of Utah. Utah State University, Logan, UT. Website http://earth.gis.usu.edu/plants/ [ac- cessed 01 September 2011]. CORBIN: A NEW ASTRAGALUS FROM UTAH 189 USDA, NRCS. 2010, The PLANTS Database. Na- tional Plant Data Center, Baton Rouge, LA.W- ebsite http://plants.used.gov [accessed 01 Oct 2010]. WELSH, S. L. 2007. North American species of Astragalus Linnaeus (Leguminosae): a taxonomic revision. Brigham Young University, Provo, UT. , N. D. ATwoop, S. GOODRICH, AND L. C. HIGGINS. 2008. A Utah flora, fourth edition, revised. Brigham Young University, Provo, UT. YONKEE, A. AND M. Lowe. 2004. Geologic map of the Ogden 7.5-minute quadrangle, Weber and Davis Coun- ties, Utah. Utah Geological Survey, Salt Lake City, UT. MADRONO, Vol. 58, No. 3, pp. 190-198, 2011 GRIMMIA VAGINULATA, (BRYOPSIDA, GRIMMIACEAE) A NEW SPECIES FROM THE CENTRAL COAST OF CALIFORNIA KENNETH KELLMAN California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118 kkellman@sbcglobal.net ABSTRACT A new California endemic species, Grimmia vaginulata, is described and illustrated. It is characterized by its very small size, julaceous habit, immersed capsule on a straight, centrally inserted seta, very large annulus, keeled unistratose leaves, autoicous sexuality, and a large cylindrical ochrea atop the vaginula that sheathes the seta to the base of the capsule. The differentiation and ecology of the new species is discussed. Key Words: California, endemic moss, Grimmia vaginulata, Grimmiaceae, new species, ochrea, vaginula. In the summer of 2008, prior to the annual meeting of the American Bryological and Li- chenological Society, Dale Vitt asked to see Orthotrichum kellmanii D. H. Norris, Shevock & Goffinet in the field. During that short trip Dale noticed tiny little plants growing on the same sandstone boulders that supported the Orthotrichum. These plants were so_ scattered and insignificant that they appeared to be immature stems of a small Grimmia, and not worth collecting. That impression proved incor- rect. Upon microscopic examination, it was immediately obvious that these plants represent- ed a remarkable plant that was unlike any other Grimmia in North America. Chief among the distinguishing characteristics was a transparent extension of the vaginula that extended to the base of the capsule. This extension, known as an ochrea, 1s not present in any North American Grimmia, and it is not mentioned in any North and Central American Grimmia literature (Flowers 1973; Crum and Anderson 1981; Crum 1994; Hastings and Greven 2007; Munoz 1999; Munoz and Allen 2002). To the casual student of the genus, this novel character suggested the possibility of a new genus. That hypothesis was supported by the scattered growth habit. A wider examination of Grimmia was undertaken resulting in the expan- sion of the North American parameters of the genus. TAXONOMY Grimmia vaginulata Kellman, sp. nov. (Figs. 1A, IC, 2, 3, 4, 5)—Type: USA, California, Santa Cruz Co., Big Basin Redwoods State Park, on dry vertical walls of calcareous sandstone boulders eroded from the sandstone bedrock in open chaparral, above the Basin Trail and below China Grade Road ca. 1.5 mi. beyond the northern intersection with SR 236, elev. 685 m, 37°12'40"N, 122°12'42’”W, 13 Aug 2008. Kellman, Vitt, & Shevock 5869 (holotype: CAS). Species Grimmia capillata De Notaris affinis, sed differt ab statura breviore, foliis statu madido vel sicco persistenter julaceis, ochrea cylindracea longiore usque ad prope basem capsulae exten- denti differt. Plants up to 2 mm tall, scattered to very loosely tufted (Fig. 5), simple, or with a short, tightly appressed branch; green when young, tan or white in age; julaceous wet or dry. Lower and perigonial leaves short ovate, 1:1, muticous, increasing in length upwards. Upper stem and perichaetial leaves (Fig. 2B) obovate or elliptical, soft when moist, weakly keeled concave, uni- stratose throughout, with at least some portion of the leaf margin hyaline, most common at the base and often extending to midleaf or slightly beyond; 1—1.65 mm long X 0.4—-0.75 mm wide (without the awn); the keel is even less pro- nounced in transverse section, appearing more convex, even tubular, especially on non-sporo- phytic plants (Fig. 4); apex acute or occasionally slightly acuminate. Basal juxtacostal cells short rectangular with thin to moderately thick, straight; relatively uniform across the base, but outer basal cells often somewhat narrower in | to 3 marginal rows. Distal and medial cells decreas- ing in length gradually from the base to the apex, 1—1.5(2.5):1, 12-16(26) um long X 7—13 um wide; rectangular, triangular, or irregularly polygonal, thick-walled, often with some portion of the lumen rounded creating small trigones, not or weakly sinuose; in transverse section plane to slightly bulging. Margins weakly recurved on one or both sides; unistratose. Costa narrow, to 40 um wide at the base, broadening toward the apex; excurrent in a hyaline, weakly toothed awn shorter than the lamina, decurrent at most 1—2 cells down the margin, and those often projecting 2011] KELLMAN: A NEW SPECIES OF GRIMMIA 19] Fic. 1. A. Grimmia vaginulata with leaves stripped exposing the sporophyte. (O) ochrea (V) entire vaginula. Scale bar = 500 um (from the type); B. G. capillata with leaves stripped exposing the sporophyte. (O) ochrea. Scale bar = 500 um (Handel-Mazetti 1778, FH: syntype of G. mesopotamica); C. Detail of typical upper leaf apex of G. vaginulata Scale bar = 80 um; D. Detail of typical upper leaf apex of G. capillata Scale bar = 175 um. as short, blunt teeth; costa in transverse section at midleaf with two homogenous rows of cells, 2 cells wide adaxially. Gonioautoicous or cladau- toicous. Calyptra irregularly crenate at the base, conical or campanulate, hyaline at the extreme -apex and at the base, naked and smooth, just covering the operculum. Vaginula (Fig. 1A), measured with ochrea, 0.5—0.6 mm long, epider- mal cells irregular, about 2—3:1, with very thin, straight walls; distally with a cylindrical ochrea surrounding but not connate to the seta and flaring just below the base of the capsule, 340— 430 um long. Seta straight, attached to the center of the capsule, 0.5 mm long. Capsule immersed 192 MADRONO G. capilla ta G. vaginulata Fic, 2. A. Sporophytic plants of Grimmia vaginulata and G. capillata showing size difference. Marks are | mm apart; B. G. vaginulata, two upper and perichaetial leaves. Scale bar = 500 um. with only the operculum exposed, irregularly wrinkled when dry, slightly wrinkled when wet, obloid, 0.9 mm long X 0.6 mm wide, abruptly contracted to the seta. Annulus (Fig. 3) of 3 rows of differentiated, transparent, thick-walled en- larged and elongated cells; remaining on the urn after dehiscence, but gradually falling off in sections. Operculum mammilose to low conical, crenulate to erose at the base. Exothecial cells irregularly rectangular to hexagonal, thin-walled, almost transparent when mature, easily revealing the stalked theca within. Stomata present. Peri- stome (Fig. 3) of 16 orange-red cribrose-dissected teeth, irregularly divided nearly to the base into 3-4 strongly spiculose filaments, ca. 185 um long. Spores smooth, 10—13 um in diameter. The specific epithet refers to the persistent cylindrical ochrea atop the vaginula, which extends nearly to the base of the capsule. IDENTIFICATION AND TAXONOMIC RELATIONSHIPS Grimmia vaginulata can be distinguished from all other North American Grimmia by the following combination of characters: very small plants that are julaceous wet or dry, upper leaves with hyaline margins, dissected peristome, and a conspicuous and persistent ochrea sheathing the entire seta to just below the base of the capsule. Grimmia vaginulata resembles G. anodon Bruch & Schimp. and G. plagiopodia Hedw. in leaf shape. However, G. vaginulata remains julaceous wet or dry, and is a much smaller and narrower plant. The straight, centrally inserted seta of G. vaginulata contrasts with the sigmoid, eccentri- cally inserted seta that characterizes both G. anodon and G. plagiopodia. The extremely dis- sected peristome resembles G. orbicularis Bruch and G. moxleyi R.S. Williams. Grimmia orbicu- laris shares the unistratose margins, and the autoicous sexuality with G. vaginulata, but the awns of G. orbicularis are evenly distributed along the stem. Grimmia moxleyi has awns restricted to the upper leaves like G. vaginulata, but the leaf margins are bistratose. Furthermore, both G. orbicularis and G. moxleyi have exserted capsules on arcuate setae. Again, both are much larger than G. vaginulata. Both gametophytically, and sporophytically, G. vaginulata appears most closely related to G. capillata De Not., a species scattered around the Mediterranean Sea (Greven 2003). They share the keeled, entirely unistratose lamina with the margins at least somewhat recurved on one or > two sides, a costa that broadens in the distal half | of the leaf, perichaetial leaves with proximal hyaline areas, and autoicous sexuality. Sporophy- tically, the two taxa are very close. Lastly, both prefer calcareous substrates. Again, G. vaginulata 2011] FIG. 3. is amuch smaller plant that stays tightly julaceous wet or dry. My examination of 16 specimens of G. capillata and G. mesopotamica Schiffn. (a syno- nym of G. capillata fide Munoz and Pando 2000; Greven 2003) from FU, MUB, and NY show plant size in G. capillata averages around 7 mm, dwarfing G. vaginulata. (Fig. 2A) The dry leaves of G. capillata are erect to erect-patent and are usually strongly keeled. Thus the costae project from the rest of the leaves giving the dry plants a more tumid and textured appearance. When moist, the leaves of G. capillata are at least erect-patent, and are relatively easy to dissect from the stem. The tiny and tightly appressed leaves of G. vaginulata are very difficult to strip for examination without tearing. Leaf size is another point of separation, with the upper stem leaves of G. capillata measuring 2—-3 mm long (Cortini Pedrotti 2001; Ignatova and Munoz 2004), while the much smaller G. vaginulata has leaves with a maximum length of 1.65 mm. In G. capillata, the more or less rounded apex (Fig. 1D) is the most commonly hyaline portion of the leaf, while in G. vaginulata, the margins are hyaline, and the apex is acute. There is some small portion at the base of the awn that is hyaline, but at most it is one to two cells down the margin (Fig. 1C). KELLMAN: A NEW SPECIES OF GRIMMIA 193 Detail of annulus and peristome of Grimmia vaginulata. Scale bar = 120 um. Fic. 4. Drawings of leaf cross sections, Grimmia vaginulata; a) leaf from sterile plant taken at midleaf; b, c, d) upper leaf from sporophytic plant. Scale bar = 100 um. 194 MADRONO FIG. 5. Although evidence is admittedly sparse, with the only known population being the type, the growth habit of G. vaginulata adds another basis for separation. In G. vaginulata, the plants are at best loosely associated (Fig. 5), not forming a contin- uous turf or clump. In G. capillata, the plants form dense green mats of sterile plants inter- spersed with a few fertile plants (Cortini Pedrotti 2001; Greven 2003; Heyn and Herrnstadt 2004; Ignatova and Munoz 2004). The base of the capsule in G. capillata is less abruptly contracted to the seta than is seen in G. vaginulata. Lastly, in G. capillata the ochrea, while pronounced, extends only half way up the seta, not to the base of the capsule as in G. vaginulata. (Figs. 1A, B) Grimmia pseudoanodon Deguchi, described in 1987 from Peru (Deguchi 1987), is another small autoicous plant (ca 5 mm tall) with immersed sporophytes with a centrally attached, straight seta, and like G. vaginulata, the base of its capsule is abruptly contracted. Examination of an isotype from NY, showed several important differences from G. vaginulata. First, G. pseudoanodon grows in cushions, contrasting the scattered habit of G. Grimmia vaginulata. Photograph of growth habit on sandstone. vaginulata. G. pseudoanodon has no peristome, its cauline leaves are not reduced down the stem, being linear to lanceolate and bearing awns to the base of the plant. They are patent to spreading when moist, and are easy to separate from the stem. Similar to G. capillata, the apex of the leaf» is hyaline for 15-20 cells, and the awn is often \ decurrent down the margins of the leaf. Most | leaves of G. pseudoanodon show bistratose | margins. The vaginula of G. pseudoanodon 1s extended, demonstrated by the aborted archego- — nia scattered throughout its length, however, the ochrea is at most represented by a few hyaline flaps at the apex of the vaginula, far from the tube sheathing the seta of G. vaginulata. Inter- | estingly, slight pressure expels the seta from the | end of the vaginula, leaving a conical stub at the | base of the seta. GENERIC TAXONOMY It is clear that this new plant belongs in the Grimmiaceae. The only other familial possibility is Ptychomitriaceae, and that is ruled out by the 2011] unlobed calyptra. Within Grimmuiuaceae, other genera can easily be dismissed as well. The large annulus, and the operculum falling independent of the columnella deny placement in Schistidium. Although the cribrose peristome resembles those of both Coscinodon and Jaffueliobryum, and the autoicous sexuality, unistratose lamina and large annulus further support placement in Jaffuelio- bryum, both genera require a pleated and somewhat sheathing calyptra (Churchill 1987; Hastings 2007; Spence 2007). The calyptra of G. vaginulata is so short that it does not even split or tear during capsule development, sitting instead just atop the operculum. Racomitrium (sensu lato) is ruled out by the lack of sinuosity in the cell walls of the lamina or the vaginula (Deguchi 1978). Not only can we suggest placement in Grimmia by elimination, but the dramatic characters displayed by G. vaginulata fit well within the limits of variation of plants already in that genus. Deguchi (1978) describes the “‘Affinis type” annulus as “‘well differentiated, composed of (2) 3-4 rows of cells, which are thick walled, but transparent, and becoming increasingly larger from the lower to the upper rows. ... Upper rows of cells of the annulus are also removed when the lid falls, but their lower rows usually remain attached to the orifice of the urn, disjoining little by little in the course of time.” This is a perfect description of the annulus in G. vaginulata. He attributes this “‘Affinis type’ annulus to “G. affinis, G. anomala, G. apiculata, G. brachydic- tyon, G. curvata, G. olympica, and G. pilifera.” Munoz (1999) assigns this annulus type to “G. involucrata, G. longirostris, G. poecilostoma, and G. trichophylla’ in the introductory section, and 14 other taxa in the species descriptions. The ochrea is a structure that is poorly understood and has received little taxonomic discussion. Deguchi (1978) makes it clear that the ochrea is only a section of the vaginula, but Magill (1990) defines ochrea as “‘vaginula, or upper part of vaginula’’. The haploid vaginula includes cells originating from both the archego- nium and the upper part of the stem (Deguchi 1978). Deguchi’s proof of this contention is the “occurrence of aborted archegonia at the basal part of the vaginule” (Fig. 1A). Maier (2002) discusses the vaginula and the ochrea and describes Grimmia donniana Sm. ex Spruce with ‘“‘ochrea broad’’, with most other Species with ochrea ‘“‘short” or “‘small’”’. Some species such as G. orbicularis (a close relative of the aforementioned G. moxleyi) she describes with ‘“‘vaginula 0.8 mm, with ochrea.” Unfortu- nately, she does not provide illustrations to inform that aspect of her excellent descriptions, nor does she describe how she measures the vaginula. Delgadillo (Universidad Nacional Autonoma de Mexico, personal communication) KELLMAN: A NEW SPECIES OF GRIMMIA Is wrote “I believe the ochrea is a fairly common feature of the neotropical species of Grimmia. It can be demonstrated in G. americana, G. donniana and G. elongata. It is particularly evident in G. anodon.”’ But more convincing examples can be found in the persistent ochreae of G. capillata and G. involucrata Cardot. Being clear then that this new plant properly belongs in Grimmia, it is necessary to at least explore its relationships within the genus. The history of the genus Grimmia, s./, 1s a tortured and complex story of confused concepts, alternating periods of splitting and lumping, and regional authors describing numerous plants that have been later synonymized in worldwide treatments (Munoz and Pando 2000; Greven 2003). Deguchi (1978) and Ochyra et al. (2003) both provide a good summary of this nomenclaturally difficult subject. Although Hernandez-Maqueda et al. (2008) casts some doubt on Coscinodon, most modern authorities agree on the separation of Jaffueliobryum, Schistidium, and Coscinodon from Grimmia, but there is disagreement whether Grimmia itself requires further subdivision. Ochyra et al. (2003) proposed to split Grimmia into five genera, with each genus corresponding in concept with the subgenera reluctantly suggested by Hastings and Greven (2007). Both systems are rooted in the work of Hagen (1909) and are summarized in Table 1. Grimmia vaginulata can be ruled out of Gasterogrimmia based on the straight, centrally attached seta. Litoneuron requires 2-stratose, concave leaves with the costa not prominently projecting abaxially: Grimmia vaginulata meets none of these criteria. An expanded Guembelia that included unistratose leaves would fit all characters of G. vaginulata, but if one were to accept the Ochyra et al. (2003) concept of Dryptodon, which includes plants with short straight setae and plicate or wrinkled capsules, then G. vaginulata would belong there. In fact, Ochyra et al. (2003) places G. capillata in Dryptodon. The difficulty is compounded, however, by statements by various authors suggesting that pairs of plants within Grimmia s./. are closely related, but subsequent authors treat each member of the pair in separate subgenera or genera. For example, G. capillata is often paired with G. crinita Brid. (Greven 2003; Ignatova and Munoz 2004) and in fact De Notaris (1836, 1838), who first described G. capillata in 1836, reduced it to a variety of G. crinita in 1838, and Maier (2010) reduced G. capillata into synonymy with G. crinita. In this pair, G. crinita is universally placed in Gasterogrimmia (Grimmia sensu Ochyra et al. 2003) but as stated above G. capillata is placed in Dryptodon. These cross subgeneric or generic affinities, as well as the taxa that defy placement within any system, illustrate the 196 TABLE 1. Hastings and Greven (2007) Gasterogrimmia Guembelia Litoneuron Rhabdogrimmia MADRONO Ochyra et al. (2003) Grimmia Orthogrimmia Guembelia Dryptodon [Vol. 58 COMPARISON OF TAXONOMIC TREATMENTS OF GRIMMIA. Description Capsules immersed, smooth, seta sigmoid and eccentrically attached to the capsule, leaves 1- or 2-stratose that are concave or concave-keeled. Capsules immersed to exserted, smooth, seta straight and centrally attached to the capsule, leaves mostly 2-stratose and keeled. Capsules exserted or emergent, smooth, seta straight and centrally attached to the capsule, leaves 2- stratose that are concave with the costa not prominent dorsally. Capsules emergent to exserted, ribbed, seta arcuate and centrally attached to the capsule, leaves mostly 1- stratose (2-stratose at the margins) and keeled. complexity of Grimmia s./. It is likely that this will only be sorted out by a massive study combining worldwide genetic and morphological data. The very dramatic and persistent ochrea in G. vaginulata opens the question of the importance of the ochrea in Grimmia taxonomy. Only Maier (2002) and Deguchi (1978) discuss this character. Deguchi briefly mentions the ochrea in his introduction, and then does not use it in his species descriptions. It is ignored in Crum (1994), Munoz (1999), Cortini Pedrotti (2001), Greven (2003), Ignatova and Munoz (2004), and Hastings and Greven (2007). Maier (2002) lists G. donniana with ochrea “‘broad’’, and G. elatior with ochrea “distinct”. Many other species were described with ochrea “‘small”’ or the character was ignored altogether. A quick examination of five specimens in CAS annotated by R. Hastings as G. donniana displayed three with no ochrea, one very juvenile capsule with an ochrea already disintegrating, and one specimen with a persistent ochrea about as long as broad at the base of a mature sporophyte. Thus, it seems that this character by itself does not seem to be reliable enough to be useful for separating G. donniana, one of the few taxa that Maier thought had a distinctive ochrea; it is likely, therefore, that the character is not useful beyond the G. capillata—vaginulata group. Nonetheless, further study of such taxa as G. involucrata may show that the ochrea could be an additional character useful for identification. ECOLOGY AND DISTRIBUTION Grimmia vaginulata has thus far only been found on vertical or underhanging surfaces on calcareous sandstone boulders that have eroded out of less calcareous sandstone bedrock of the Butano Formation. These rocks are scattered through a variably dense chaparral comprised of Adenostoma fasciculatum Hook. & Arn., Arcto- staphylos tomentosa (Pursh.) Lindl., Ceanothus cuneatus (Hook.) Nutt., C. papillosus Torr. & A. Gray, and Eriodictyon californicum Hook. & Arn.. The climate is Mediterranean with moder- ate but rainy winters, and hot rainless summers (Kellman 2003). The elevation is slightly below 700 meters, placing the site above all but the thickest summer maritime fog. The lack of summer fog keeps these sites very hot and dry until the winter storms arrive. One of the more peculiar aspects of this very interesting plant is the substrate. With the exception of one recent collection from marble rock (Kellman, Shevock & Lodder 6133 [CAS]) the boulders are also the sole known substrate of © Orthotrichum kellmanii, another rare coastal California endemic (Norris et al. 2004). The bedrock was deposited in the Upper Eocene, and is part of the Butano Sandstone Formation (Brabb 1989), but the geologic history of the boulders is not known. Presumably, the rocks were formed and eroded into their rounded shape some time prior to the Eocene deposits that | formed the bedrock. In Santa Cruz Co., the | Butano Formation is exposed along the western slopes of the highest mountains on the eastern © side of the county (Brabb 1989), but the boulders are only present in a small patch in Big Basin | Redwoods State Park. There is also a small field | of the boulders in southern San Mateo Co., also | in Butano State Park. Recently, this same | formation with exposed boulders has been | found in Monterey Co. within the Ventana | Wilderness of the Los Padres National Forest. | Orthotrichum kellmanii was collected here al- | though much of the area remains to be surveyed. The new population greatly expanded its range | to the south. Although access to these sandstone | outcrops was difficult due to hiking through 2011] dense stands of chaparral, the area subsequently burned in the summer of 2008. This area 1s now among the highest priority sites to survey for G. vaginulata—and to document the extent of Ortho- trichum kellmanii in this area. Associated bryophytes growing on and around the boulders include Amphidium californicum (Hampe ex Mill. Hal.) Broth., Antitrichia cali- fornica Sull. in Lesq., Gemmabryum californicum (Sull.) Spence, Didymodon vinealis (Brid.) R.H. Zander, Tortula muralis Hedw., Grimmia torenti Hastings, G. pulvinata (Hedw.) J. E. Smith, Gymnostomum calcareum Nees & Hornsch., G. viridulum Brid., Orthotrichum kellmanii, and Cephaloziella divaricata (Sm.) Warnst. CONSERVATION At the type location, G. vaginulata has been found on only four out of several hundred boulders, and these fit within a circle of ten meters. It is imperative that no further collections should be made until more populations are found. Additional surveys need to be conducted throughout the highly restricted habitat within the Butano Formation within State Park Lands and a federally designated wilderness area. Even if additional occurrences are discovered, Grimmia vaginulata will almost certainly remain a very narrowly restricted endemic that could be ad- versely impacted by stochastic events. Even though the chaparral is a fire adapted ecosystem (Schoenherr 1992), the extreme rarity of G. vaginulata leaves open the possibility of fire killing the entire known population. In March of 2009, the author visited the Bonny Doon Ecolog- ical Preserve in Santa Cruz Co. Nine months before, in June of 2008, a very hot fire raced through the preserve, where sandstone rocks are scattered through a chaparral very similar to that found at the type location of G. vaginulata. Even in places where the brush was minimal, virtually all bryophytes were killed on the rocks, especially those bryophytes growing on the walls and underhanging surfaces of the rocks. A few mosses and liverworts survived on the tops of a few rocks. Apparently the heat of the fire was trapped under the rocks and then bathed the vertical surfaces, killing all plant life. It is conceivable that thick cushions, or gemmiform plants with densely imbricate leaves could insulate at least some part of a dry moss plant from the heat, but the loose colonies and cylindrical plant form of G. vaginu- lata offers no such protection. To date, G. vaginulata is known only from the type and in general, taxonomic novelties should not be based on a single specimen, particularly in variable genera such as Grimmia. However, this plant’s combination of characters, unique to Grimmia in North America, along with the conservation implications of its rarity, demand a KELLMAN: A NEW SPECIES OF GRIMMIA LOT full description. Of course, it is always possible that further collections from new locations could alter the concept of G. vaginulata. Of all the characters discussed above, the most likely features to change would be the size and the growth form. It is possible that other populations of G.vaginulata could be composed of larger plants. Arguing against that possibility is the fact that both sterile and fruiting plants exhibit the same size and growth form in the type population. It seems more likely that G. vaginulata could be found growing in turfs or small cushions. However, turf and cushion formation require the survival of the originating stems. There are no subapical innovations in the type population which suggest that this species may be short lived and that individuals die after sexual reproduction. ACKNOWLEDGMENTS I am indebted to Jim Shevock and Dan Norris for teaching me how to do the research necessary to write a paper of this type. They may or may not have been aware that they were teaching me these methods, but an interested student learns as much by observation as by lecture and discussion. I honor their example. I would also like to thank Roxanne Hastings for teaching me the basic understanding of Grimmia, and for a thorough review of this paper resulting in many positive changes. I thank Henk Greven for his review of this paper, and the suggestion to compare Grimmia vaginulata with G. pseudoanodon. David Toren deserves credit for pointing out the illustration of Grimmia mesopotamica in the Bryophyte Flora of Israel that put me onto the similarities of G. vaginulata and G. capillata. Thanks also to Eva Maier, and Claudio Delgadillo, for their insights on the vaginula and ochrea. I also appreciate Claudio Delgadillo and Jesus Munoz for their helpful comments on an earlier version of this paper. Steve Lodder was invaluable in providing the beautiful photographs. I thank Patricia Eckel for providing the Latin description. I would like to thank the curators of MUB, FH, and NY for the loan of specimens of Grimmia capillata, Grimmia mesopotamica, and Grimmia pseudoanodon, and Deb Trock at CAS who arranged for the loans. LITERATURE CITED BRABB, E. E. 1989. Geologic map of Santa Cruz County, California. Miscellaneous Investigation Series. U.S. Geological Survey, Denver, CO. CHURCHILL, S. P. 1987. Systematic and biogeography of Jaffueliobryum (Grimmiaceae). Memoirs of the New York Botanical Garden 45:691—708. CORTINI PEDROTTI, C. 2001. Flora dei muschi d’Italia: Sphagnopsida, Andreaeopsida, Bryopsida (1 Parte). Antonio Delfino Editore, Rome, Italy. CruM, H. 1994. Grimmiales. 386-415. in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mexico. Memoirs of the New York Botanical Garden 69:1—1113. AND L. ANDERSON. 1981. Mosses of Eastern North America. Columbia University Press, New York, NY. DE NOTARIS, G. 1836. Memorie della Reale Accademia delle Scienze di Torino 39:248. 198 MADRONO . 1838. Syllabus Muscorum in Italia et in Insulis Circumstantibus Hucusque Cognitorum, Turin, Italy. DEGUCHI, H. 1978. A revision of the genera Grimmia, Schistidium and Coscinodon (Musci) of Japan. Journal of Science of the Hiroshima University Series B, Div. 2 (Botany) 16:121—256. . 1987. Studies on some Peruvian species of the Grimmiaceae. Pp. 19-74 in H. Inoue (ed.), Studies on cryptogams in Southern Peru. Tokai University Press, Tokyo, Japan. FLOWERS, S. 1973. Mosses: Utah and the West. Brigham Young University Press, Provo, UT. GREVEN, H. C. 2003. Grimmias of the world. Bachuys, Leyden, The Netherlands. HAGEN, I. 1909. Forarbejder tilen Norsk lovmosflora. IX. Grimmiaceae. Det Kongelige Norske Videns- kabers Selskabs Skrifter 1909(5):1—94. HASTINGS, R. I. 2007. Coscinodon. Pp. 258-262 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, Vol. 27. Oxford University Press, New York, NY , H. C. GREVEN. 2007. Grimmia. Pp. 225-258 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, Vol. 27. Oxford University Press, New York, NY. HERNANDEZ-MAQUEDA, R., D. QUANDT, O. WERNER, AND J. MUNOZ. 2008. Phylogeny and classification of the Grimmiaceae/Ptychomitriaceae complex (Bryophyta) inferred from cpDNA. Molecular Phylogenetics and Evolution 46:863—877. HEYN, C. C. AND I. HERRNSTADT. 2004. The bryophyte flora of Israel and adjacent regions. The Israel Academy of Sciences and Humanities, Jerusalem, Israel. IGNATOVA, E. A. AND J. MUNOZ. 2004. The genus Grimmia in Russia. Arctoa 13:101—182. [Vol. 58 KELLMAN, K. M. 2003. A catalog of the mosses of Santa Cruz County, California. Madrofio 50:61-82. MAGILL, R. E. 1990. Glossarium Polyglottum Bryolo- giae, a multilingual glossary for bryology. Missouri Botanical Garden, Saint Louis, MO. MAIER, E. 2002. The genus Grimmia (Musci, Grimmia- ceae) in the Himalaya. Candollea 57:143—238. . 2010. The genus Grimmia Hedw. (Grimmia- ceae, Bryophyta) - a morphological-anatomical study. Boissiera 63:1—377. MuNoz, J. 1999. A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Annals of the Missouri Botanical Garden 86:118—191. AND F. PANDO. 2000. A world synopsis of the genus Grimmia (Musci, Grimmiaceae). Missouri Botanical Garden Press, St. Louis, MO. AND B. ALLEN. 2002. Grimmia. Pp. 221—231 in B. Allen (ed.), Moss flora of Central America, Part 2. Encalyptaceae—Orthotrichaceae. Missouri Botanical Garden Press, Saint Louis, MO. Norris, D. H., J. R. SHEVOCK, AND B. GOFFINET. 2004. Orthotrichum kellmanii (Bryopsida, Ortho- trichaceae), a remarkable new species from the central coast of California. The Bryologist 107:209-214. _ OCHYRA, R., J. ZARNOWIEC, AND H. BEDNAREK- OCHYRA. 2003. Census catalog of Polish mosses. Polish Academy of Sciences, Krakow, Poland. SCHOENHERR, A. A. 1992. A natural history of California. University of California Press, Berkeley and Los Angeles, CA. SPENCE, J. R. 2007. Jaffueliobryum. Pp. 262-264 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, Vol. 27. Oxford University Press, New York, NY. MADRONO, Vol. 58, No. 3, pp. 199-200, 2011 REVIEW Introduction to California Chaparral. By RONALD D. QUINN AND STERLING C. KEELEY. 2006. University of California Press, Berkeley, CA. 344 pp. ISBN 9780520219731, $55.00, hardcover; ISBN 9780520245662, $21.95, paperback. In Introduction to California Chaparral, au- thors Ronald D. Quinn and Sterling C. Keeley condense the beauty and diversity of California’s most prominent habitat into a beginners guide suitable for curling up with on the couch or toting along on a hike. The diminutive tome attempts to familiarize the casual observer not only with that superficial ““‘bluish-green blanket gently covering the hills,’ but also with the dynamic interplay of organisms and underlying ecological processes beneath that oft go largely unseen. It is a monu- mental task to be sure. To accomplish this task, the authors guide us through the climatic forces that shape the system; the dominant plant, animal and insect species that inhabit it; and other forces such as fire, flooding and urban encroachment that affect it. In addition to describing the chaparral world that modern day adventurers might observe, Quinn and Keeley take us briefly through the history of the chaparral to provide a backdrop of the forces that created such diversity. In distilling such broad and complex subject matter, the authors only stumble in perhaps the most complex of chaparral arenas: wildfire. Dedicating roughly 1/6 of the book specifically to fire and regularly referring to it throughout the rest, the authors succeed in divulging most of the issues surrounding fire in chaparral. However, their attempt at presenting those issues together in a cohesive and coherent whole falls short and proves dangerously misleading. Notwithstanding, the authors bring hundreds of years of scientific discovery, field observation and a love of Cali- fornia’s defining ecosystem into an accessible and enjoyable guide for the masses. This attractive collection consists of 302 pages of text, color photos, black and white diagrams, plus a glossary and reference section for those who wish to dig deeper. All this is printed on thick, sturdy pages that will no doubt survive the jostles of the many hikes it is sure to inspire. Quinn and Keeley, Professors of biological science and botany, respectively, draw on a wide range of sources and personal research in com- piling this guide. They use these sources to paint a colorful mural of wood-rat nests, heat-seeking fire beetles, iridescent hummingbirds and an impressive array of plants that have managed to survive eons of yearly drought and occasional wildfire. The descriptions are presented at an accessible level of detail for its intended audience. Plant and animal common names are provided in conjunction with their scientific equivalents. Throughout the guide, the particular adaptations that help chaparral plants and animals survive this difficult environment, 1.e., the very features that make them uniquely chaparralian, are highlighted. Reference especially is often given to a particular species’ relationship to fire. The authors provide not only colorful descriptions but interesting facts that keep even the most hardened urbanite engaged. Unfortunately, colorful language occasionally gives way to literary excess. The authors too often employ personification, especially with respect to plant characteristics, that can undermine the integrity of the data presented. An example on the innocuous end of the spectrum involves a species of monkey flower, Mimulus aurantiacus. The authors describe the unique stigma that assumes a closed position when pollinated or even just touched by an insect or animal. The authors then claim that in so doing the plants ‘“‘advertise their status” as pollinated or not, presumably giving pollinators the plant kingdom’s equivalent of a wedding ring to would- be suitors. Does a closed stigma make it more likely that pollinators will move to another flower, or is there some other evolutionary incentive for the stigma closing? It’s a question the authors don’t address, nor is the conclusion supported with cited research. In this case, it’s an admittedly harmless conclusion. The authors’ phrasing becomes critically dangerous with regard to fire. The first example comes in the punctuating sentence of the introductory paragraphs to the fire section. It reads: “In short, where there is chaparral, there is fire.” While this may technically be true, it is misleading. Where there is chaparral there is going to be fire. But where there are grasslands there is fire. Where there are conifer forests there is fire. In other words, the presence of fire in chaparral is not due to something inherent in the plant community. Rather, where there is chapar- ral there is a Mediterranean climate and where there is a Mediterranean climate (or any other dry climate with enough moisture to sustain vegetative growth) and an ignition source there will be fire, regardless of vegetation type. The phrase implies that chaparral fosters a fire environment rather than the more accurate paradigm that chaparral species exist because they are able to survive infrequent fires. The result of the authors’ approach is a_ subtle perpetuation of the oft repeated and poorly 200 applied concept that chaparral needs to burn. As the authors themselves state, certain species do produce seeds that require fire cues to germinate. On the other hand, we know chaparral stands existing for more than a century without fire are perfectly healthy systems and if chaparral stands burn too frequently they can be converted to non-native grasslands. Extrapolating this dichot- omy is a tightrope that the authors have difficulty walking. The most prominent example comes in their ‘Living with the Chaparral” chapter. In it, the authors describe the merits of prescribed fire and clearing fuel breaks, offering them as effective means to reduce mature chaparral and thus reduce the hazards fire presents to humans. Young chaparral, they suggest, burns less intense- ly and cuts off the fuel supply, extinguishing fires. Later, not as a counterpoint but as a discrete section, they explain that extreme weather condi- tions drive fires right through fuel breaks and fuel-age mosaics presumably negating the value of such “‘barriers”’. Further on, the intrinsic watershed value of mature chaparral stands is described in detail. The authors explain that ““mature chaparral acts as a sponge”’ retaining rainfall in the soil and aquifers beneath. Without a mature chaparral and root system, water skims off the soil surface creating disastrous debris flows and leaving no water behind for the ecosystem. That the previously advocated prescribed burning and man-made fuel breaks destroy this important ““sponge’’ mecha- nism is a conclusion never fully reconciled by the authors. In truth, land management with respect to wildfire, watershed values, and protecting lives and property is very much a balancing act that few, if any, have yet to master. It is perhaps unfair MADRONO [Vol. 58 then to expect the authors to provide solutions to this dilemma. Rather than offer a cohesive recommendation, the authors drop all the issues in our lap and leave us to sort them out. It is a safe approach. Nevertheless, since this work appears to imply that mature chaparral is a threat to humans rather than the other way around, it will likely draw ire from some ecologists and conser- vationists. And in that sense it is perplexing to find the text demonizing an ecosystem because it can be inconvenient to humans. While perhaps not packaged in the most ideal manner, the authors at least conclude the wildfire section with a plea for humans to take responsibility for their urban planning practices to avoid further losses. Muddled fire issues aside, this 1s an ““/ntroduc- tion to California Chaparral” and in that sense it serves its purpose well. Quinn and Keeley’s work is a celebration of a unique and teeming habitat. To the reader unfamiliar with that amorphous blanket of shrubs covering the hillsides outside their car windows, Introduction to California Chaparral provides a comprehensive glimpse into that complex habitat. Where the authors partic- ularly shine is in capturing the sheer diversity and dynamic drama found within the often impene- trable chaparral canopy. Were the reader to carry this guide out into the field with them they would likely recognize many more twitters, rustles and floral displays than ever before. Ultimately, creating that connection to the natural environ- ment is the greatest gift any environmental author can bestow upon a reader. In Introduction to California Chaparral Quinn and Keeley are likely to succeed in doing so many times over. —CHRISTOPHER BLAYLOCK, California Chaparral Insti- tute, P.O. Box 545, Escondido, CA 92033; chris@ blaylock.org. MADRONO, Vol. 58, No. 3, p. 201, 2011 NOTEWORTHY COLLECTION CALIFORNIA PORPHYRA SUBORBICULATA Kjellman_ 1897:10—13 (BANGIACEAE).—Marin Co., epizoic on barnacles attached to pilings in the upper intertidal at Nick’s Cove, Tomales Bay, 38°11'57.42”N, 122°55'16.40"W, thalli sterile and fertile, 12 May 2011, J. R. Hughey s.n. (UC 1966687, UC 1966688) and 20 July 2011, J. R. Hughey s.n. (UC 1966689). Previous knowledge. Porphyra suborbiculata is natu- rally distributed throughout Asia (type locality: Goto- retto, Nagasaki Prefecture, Japan (Silva et al. 1996) and reportedly also occurs naturally in the Indian Ocean, Australia, and New Zealand (Broom et al. 2002). However, based on recent molecular evidence some P. suborbiculata from Australia and New Zealand, as well as North America (Mexico and the Atlantic coast of North America) represent introduced populations (Broom et al. 2002; Klein et al. 2003; Neefus et al. 2008). The occurrence of this seaweed in areas associated with increased shipping activity suggest that its numerous introductions were the result of hitchhik- ings on the hulls of seagoing vessels (Broom et al. 2002). In the eastern Pacific, P. suborbiculata was first collected in 1985 from Baja California, Mexico (Aguilar-Rosas and Aguilar-Rosas 2003). Although P. suborbiculata shows some variation in morphology, it is generally characterized as being a relatively small (0.5— 4 cm in diameter), monostromatic alga, with brownish red to pink, or bronze and violet colored blades that are ovate to cordate or reniform in shape (Broom et al. 2002; Aguilar-Rosas and Aguilar-Rosas 2003; Neefus et al. 2008). Significance. First report of P. suborbiculata in California. The gametophytic thalli collected from Tomales Bay are in agreement with descriptions and illustrations of this species. Specimens from California are cordate at the base, and ovate to deeply reniform in shape. The margins are slightly ruffled and appear dentate to the unaided eye. Under microscopic exam- ination thalli show the diagnostic marginal teeth. The blade color is reddish-brown in spring and steel greenish in summer, and fronds are more or less equal in width and height, measuring to 1.8 cm. Spot checks for this alga at Marshall and Marconi failed to yield additional specimens. Identification of this invasive species was confirmed using rbcL (GenBank JN413680) and ITS-1 (GenBank JN413679) DNA _ sequences. The rbcL sequence was identical to two sequences from Japan (Kanagawa, Yokosuka, Sajima and Yamaguchi, Shi- monoseki, Tsunoshima) and differed from others in the database by 2 or more bp. The ITS-1 sequence was identical to fifteen other sequences deposited in GenBank representing populations from around the world. Since the rbcL sequence generated for P. suborbiculata from Tomales Bay matches specimens from Japan and the ITS-1 sequence is identical to other invasive populations of this species from Australia, Baja California, New Zealand, and the western Atlantic, it is concluded that the population from California is also the result of an introductory event. In Tomales Bay, P. suborbiculata joins a list of five other non-native algae: Caulacanthus ustulatus (Turner) Kutzing, Codium fragile subsp. tomentosoides (van Goor) P. C. Silva, Gelidium vagum Okamura, Lomentaria hakodatensis Yendo, and Sargassum muticum (Yendo) Fensholt (C. K. Kjeldsen, Sonoma State University, unpublished data; Hughey 1995; Hughey et al. 1996). The maricul- ture of oysters in Tomales Bay began around 1907 (Barrett 1963), and oysters are the likely vector for the introduction of P. suborbiculata in the bay. —JEFFERY R. HUGHEY, Department of Science and Engineering, Hartnell College, Salinas, CA 93901. jhughey@hartnell.edu. LITERATURE CITED AGUILAR-ROSAS, R. AND L. E. AGUILAR-ROSAS. 2003. El género Porphyra (Bangiales, Rhodophyta) en la costa Pacifico de México. I. Porphyra suborbiculata Kjellman. Hidrobioldgica 13:51—S6. BARRETT, E. M. 1963. Fish Bulletin 123. The California oyster industry. Scripps Institution of Oceanogra- phy Library, UC San Diego, San Diego, CA, Website http://escholarship.org/uc/item/1870g57m [accessed 10 September 2011]. BRooM, J. E., W. A. NELSON, C. YARISH, W. A. JONES, R. AGUILAR ROSAS, AND L. E. AGUILAR ROSAS. 2002. A reassessment of the taxonomic status of Porphyra suborbiculata, Porphyra caroli- nensis and Porphyra lilliputiana (Bangiales, Rho- dophyta) based on molecular and morphological data. European Journal of Phycology 37:227—235. HUGHEY, J. R. 1995. A systematic study of Chondra- canthus Kutzing (Rhodophyceae) with a contribu- tion to the marine flora of Tomales Bay, California. M.A. thesis, Sonoma State University, Rohnert Park, CA. , C. K. KJELDSEN, P. C. SILVA, R. L. MOE, AND T. C. DECEW. 1996. Noteworthy Collections. Madrono 43:432. KLEIN, A. S., A. C. MATHIESON, C. D. NEEFUS, D. F. CAIN, H. A. TAYLOR, B. W. TEASDALE, A. L. WEST, E. J. HEHRE, J. BRODIE, C. YARISH, AND A. L. WALLACE. 2003. Identification of north- western Atlantic Porphyra (Bangiaceae, Bangiales) based on sequence variation in nuclear SSU and plastid rbcL genes. Phycologia 42:109—122. NEEFUS, C. D., A. C. MATHIESON, T. L. BRAY, AND C. YARISH. 2008. The distribution, morphology, and ecology of three introduced Asiatic species of Porphyra (Bangiales, Rhodophyta) in the north- western Atlantic. Journal of Phycology 44:1399— 1414. SILVA, P. C., P. W. BASSON, AND R. L. MOE. 1996. Catalogue of the benthic marine algae of the Indian Ocean. University of California Publications in Botany, Berkeley, CA. MADRONO, Vol. 58, No. 3, pp. 202-203, 2011 NOTEWORTHY COLLECTION CALIFORNIA SEQUOIADENDRON GIGANTEUM (Lindl.) J. Buchholz (CUPRESSACEAE) (giant sequoia, big tree, or Sierra redwood).—Riverside Co., northwestern San Jacinto Mts., northeast of HW 243 about 18 highway km NW of Idyllwild, in the southern unit of the San Bernardino National Forest, on the northwestern flank of Black Mt. along both sides adjacent to and well back from the Black Mountain Trail, from upper Hall Canyon, through the sloping plateau or saddle near the westerly ridge, and into the final switchback ascent to the summit (elev. 2369 m). Species seen from 2036 to 2236 m elev. in mixed-conifer forest (Lower Montane Conifer- ous Forest) on sandy granitic soil; associates include: Abies concolor, Pinus coulteri, P. ponderosa var. pacifica, and Quercus chrysolepis dominant at lower elevations; P. jeffreyi and P. lambertiana dominant at higher elevations; Calocedrus decurrens, P. flexilis, and Ribes nevadense throughout most of area; 1 May 2009. Collections (vouchers to be deposited at RSA, UC, and UCR). Tree DBH 65 cm, sterile, just left (as ascending) of trail on slope by large trunk lying across trail, N33 49.540, W116 45.977, 2036 m elev., R. Schmid & M. Schmid 2009-1; tree with abundant male cones, right (as ascending) of trail on slope covered with Pteridium aquilinum and near drainage (Indian Creek) with Alnus rhombifolia, N33 49.579, W116 45.891, 2066.5 m elev., R. Schmid & M. Schmid 2009-2; mature seed cones with opened cone scales and dislodged seeds, plus mix of sandy granitic soil and sparse duff, along Black Mountain Trail above drainage (Indian Creek) and below sloping plateau or saddle, ca. 2070 m elev., R. Schmid & M. Schmid 2009-4; tree DBH 20 cm, with medium-sized female cones, right (as ascending) side of trail in flat open area, N33 49.607, W116 45.854, 2093 m elev., R. Schmid & M. Schmid 2009-5; tree ca. 5.5 m high, with many, very large, immature female cones, just left (as ascending) of trail in flat open area, N33 49.713, W116 45.773; 2144 m elev., R. Schmid & M. Schmid 2009-6. In addition to these collections made on 1 May 2009, we did a GPS census in the vicinity of the Black Mountain Trail, starting in upper Hall Canyon. The census revealed both in the canyon and upslope beyond it at least 157 individuals from 2036 to 2236 m elev. (ca. 0.7 km linear distance, ca. 0.1 km’), plus an outlier sapling 450 m distant at 2361 m elev. near the summit of Black Mt. (2369 m). Our set of plots involved four groups at progressively higher elevations: (1) at the head of Indian Creek in the drainage (including vouchers 2009-1 and 2009-2); (2) on the slope coming out of the drainage (including voucher 2009-4); (3) at the sloping plateau or saddle (including vouchers 2009- 5 and 2009-6); (4) on the northwest-facing slope closer to the summit. This species alien to southern California is regenerating prolifically on Black Mt., as revealed by multiple age classes, from seedlings and _ saplings (juveniles) about 20-60 cm tall to trees over 6 m tall, about 40 years old, and reproductively mature. Previous knowledge. The monotypic California en- demic Sequoiadendron giganteum is native to the western slope of the Sierra Nevada, where it occurs in isolated groves in mixed-conifer forest (Lower Montane Coniferous Forest) between 825 and 2700 m elev. The 67 groves are mostly of very restricted extent and/or threatened. They occur in a narrow strip measuring about 395 km long (northwest-southeast) and 19-21 km at the widest points (east-west) and extending over seven counties from southern Placer Co. southeasterly to southern Tulare Co. (Flint 2002; Willard 2000). This species has been extensively planted in Califor- nia as an ornamental, as part of afforestation attempts, and as reforestation efforts, especially post-fire re- vegetation ones. For example, Burns and Sauer (1992) noted that 22,900 seedlings of S. giganteum were planted in southern California in the San Gabriel Mts. alone, but neither this species nor 44 other alien conifer species planted there as part of afforestation projects “‘are invading adjacent natural habitats” (p. 49). However, Rogers (1986: p. 33) wrote: “On the San Bernardino [National Forest] about 5000 to 10,000 seedlings are planted each year, and at least one instance of natural regeneration from some of the early plantings has been observed.’’ Rogers’s “‘one instance”’ may well be the introduction of S. giganteum after the Aug. 1974 fire in Hall Canyon (Keeler-Wolf 1990; Cheng 2004) and its subsequent naturalization on Black Mt., which is the subject of the present preliminary report. Our extensive analysis of the print and Internet literature for the floristics and ecology of southern California suggests S. giganteum is possibly also naturalized in the San Gabriel Mts. of Los Angeles Co. and southwestern San Bernardino Co., and in the San Bernardino Mts. of San Bernardino Co. Significance. First report and collections for River- side Co. and the San Jacinto Mts. (see the database of the Consortium of California Herbaria, http://ucjeps .berkeley.edu/consortium). More importantly, first doc- umented record for naturalization of this Sierra- Nevada endemic species in montane southern California. State and regional floras for California should acknowledge in their keys and descriptions such naturalizations. Details of these findings will be published elsewhere (Schmid and Schmid in press). —RUDOLF SCHMID, Department of Integrative Biology, University of California, Berkeley, CA 94720-3140; MENA SCHMID, Somerville, MA 02144. schmid@berkeley.edu. LITERATURE CITED BURNS, C. AND J. SAUER. 1992. Resistance by natural vegetation in the San Gabriel Mountains of California to invasion by introduced conifers. Global Ecology and Biogeography Letters 2:46—51. CHENG, S. (tech. ed). 2004. Forest Service Research Natural Areas in California. General Technical Re- port PSW-GRT-188. USDA Forest Service, Pacific Southwest Research Station, Albany, CA. Website http://www. fs.fed.us/psw/publications/documents/ psw_gtr188/ [accessed 20 June 2011]. FLINT, W. D. 2002. To find the biggest tree, revised ed. Sequoia Natural History Association, Three Riv- ers, CA. 2011] KEELER-WOLF T. 1990. Ecological surveys of Forest Service Research Natural Areas in California. Gen- eral Technical Report PSW(GTR)-125. USDA Forest Service, Pacific Southwest Research Station, Berkeley, CA. Website http://www.fs.fed.us/psw/publications/ documents/psw_gtr125/ [accessed 20 June 2011]. ROGERS, R. R. 1986. Management of giant sequoia in the national forests of the Sierra Nevada, California. Pp. 32-36 in C. P. Weatherspoon, Y. R. Iwamoto, and D. D. Piirto (tech. coordi- nators). Proceedings of the workshop on man- agement of giant sequoia, May 24-25, 1985, NOTEWORTHY COLLECTION 203 Reedley, California. General Technical Report PSW(GTR)-95. USDA Forest Service, Pacific Southwest Forest and Range Experimental Sta- tion, Berkeley, CA. Website http://www. fs.fed.us/ psw/publications/documents/psw_gtr095/_— [ac- cessed 20 June 2011]. SCHMID, R. AND M. SCHMID. In Press. Naturalization of Sequoiadendron giganteum (Cupressaceae) in montane southern California. Aliso 30. WILLARD, D. 2000. A guide to the sequoia groves of California. Yosemite Association, Yosemite Na- tional Park, CA. MADRONO, Vol. 58, No. 3, p. 204, 2011 NOTEWORTHY COLLECTION COLORADO DRABA WEBERI R.A. Price & Rollins (BRASSICA- CEAE).—Park Co., Middle Fork of the South Platte, near Magnolia Mill, Pike National Forest, in mossy depressions on north side of granite boulder, 10 plants in flower and fruit, at 39.36112°N, 106.09013°W, 3,398 m elev., 20 July 2010, Bernadette Kuhn 7927a and 7927b, along with Gina Glenne, Sheila Lamb, Alicia Langton, Ellen Mayo, Steve Olson, and Jeff Sprovkin (MO). Dr. Ihsan Al-Shehbaz verified the identification. Previous knowledge. Global distribution is limited to the type locality in the Monte Cristo Creek drainage, below the Upper Blue Lake Reservoir Dam, on the west side of Continental Divide, Summit County, Colorado. As described in the original publication (Price and Rollins 1991) the holotype (Price #464) was collected from “‘crevices of rocks beside cascades, amid rocks at edge of stream’. The type locality was described by Price as containing “‘perhaps 100 individuals”, and in 2009, 81 individuals were reported present at the site (Colorado Natural Heritage Program 2011). All indi- viduals documented are on private land owned and operated by Colorado Springs Utility, though the site is surrounded by National Forest System lands (Decker 2006). Significance. Second documented occurrence. For twenty years botanists have been searching for another occurrence outside the species’ type locality. Our collection, located 2.8 km to the south, expands the known distribution of D. weberi to another county (Park), and to the east side of the Continental Divide. In addition, it confirms the long-suspected presence of D. weberi on National Forest System land. This finding, along with the recent addition of D. weberi to the USDA Forest Service Sensitive List, may present new opportunities for the management and conservation of this extremely rare Colorado endemic (USDA Forest Service 2011). —BERNADETTE KUHN, Colorado Natural Heritage Program, 254 General Services Building, 1474 Campus Delivery, Fort Collins, CO 80523-1474. bernadette. kuhn@colostate.edu. LITERATURE CITED COLORADO NATURAL HERITAGE PROGRAM. 2011, Biodiversity Tracking and Conservation System (BIOTICS). Colorado State University, Fort Col- lins, CO. Website http://www.cnhp.colostate.edu/ exchange/request.asp [accessed 01 May 2011]. DECKER, K. 2006. Draba weberi Price & Rollins (Weber’s draba): a technical conservation assess- ment. USDA Forest Service, Rocky Mountain Region, Golden, CO. Website http://www. fs.fed.us/ r2/projects/scp/assessments/drabaweberi.pdf [ac- cessed 1 May 2011]. PRICE, R. A. AND R. C. ROLLINS. 1991. New taxa of Draba (Brassicaceae) from California, Nevada, and Colorado. Harvard Papers in Botany 3:75—77. USDA FOoRrRESsT SERVICE. 2011. Supplement No. 2600- 2011-1. Chapter 2670—Threatened, endangered, and sensitive plants and animals. USFS Forest Service Manual, FSM 2600-Wildlife, Fish, and Sensitive Plant Habitat Management. USDA Forest Service, Rocky Mountain Region (Region 2), Denver, CO. Website http://www.fs.fed.us/im/ directives/field/r2/fsm/2600/fsm_r2_2670_201 1-1. docx [6 January 2012]. MADRONO, Vol. 58, No. 3, p. 205, 2011 NOTEWORTHY COLLECTION MEXICO ANEMONE TUBEROSA Rydb. (RANUNCULA- CEAE).—Baja California, Municipio of Ensenada, Ejido Nativos del Valle heading W to Santo Tomas, 31.42561°N, 116.34858°W (WGS 84), 434 m/1428 ft, 26 March 2010, Sula Vanderplank, Sean Lahmeyer, Ben Wilder and Karen Zimmerman 100326-29 (RSA), Less than 100 plants observed growing with Zigadenus sp. on N-facing side of a steep limestone outcrop. Most plants in full flower. Previous knowledge. The general habitat of this species is given by Dutton et al. (1997), in the Flora of North America, as from rocky slopes and stream sides 800-2500 m. It is known from desert regions of the southwestern USA (CA, NM, NV, TX, UT) and NW Mexico (Baja California, Sonora), and has been well documented within the Sonoran and Mojave deserts: Wiggins (1980) reports it from the western edge of the Colorado Desert and the eastern Mojave Desert; Wilken (1993) gives the range in California as eastern Desert Mountains; 900-1900 m; Munz (1974) reports elevations of 3000—5,000 ft (914-1520 m), in Joshua Tree Woodland and Pinyon-Juniper Woodland, west- ern edge of the Colorado Desert, eastern Mojave Desert. The westernmost known collections for this taxon in California are from the southern Cuyamaca Mtns., near the southwestern end of Poser Mtn, March 19 1995, Jeri Hirshberg 253 (RSA) 60 km from the Pacific coast; and the Laguna Mountains, below Desert View, 06 April 1939, A. J. Stover 245 (SD), 80 km from the Pacific coast (CCH 2011). In Arizona, Kearney and Pebbles (1951) report elevations of 2,500—S000 ft (760— 1520 m). In Mexico, the two Baja California collections closest to the Pacific Ocean are both 50 km inland (30— 45 km from the Gulf of California to the east) in the central desert, close to the narrowest part of the peninsula: summit of volcanic hill at top of Jaraguay Grade, 24 Feb 1973, Reid Moran 20248 (SD), elev. 875 m, (the lowest documented elevation for this species in the state); Sierra San Borja, summit of Cerro la Chona, 19 Mar 1966, Reid Moran 12782 (SD), 1450 m (San Diego Natural History Museum 2011). There are a small number of California specimens from ~400 m elev., all from a small area within the Whipple Mountains, on the border with Arizona (CCH). Significance. First collection of this species west of the peninsular ranges and inside the California Floristic Province. This collection represents a range extension of ca. 70 km west from the peninsular ranges, and is the most coastal collection documented to date, being just 15 km (9 mi) from the Pacific Ocean. The elevation is the lowest recorded in Baja California, and is near the lowest elevation recorded for the species anywhere. Further investigation of the foothills surrounding this site may help determine if this small population is indeed isolated from the core range, perhaps as a result of the unusual habitat provided by the limestone outcrop. —SULA VANDERPLANK, Deptartment of Botany & Plant Sciences, University of California, Riverside, CA, 92507; SEAN LAHMEYER, The Huntington Library, Art Collections, and Botanical Gardens, 1151 Oxford Road, San Marino, CA 91108. sula.vanderplank@ gmail.com. LITERATURE CITED CONSORTIUM CALIFORNIA HERBARIA (CCH). 2011. University of California, Berkeley, CA, Website http://ucjeps.berkeley.edu/consortium/ [accessed 3 March 2011]. DUTTON, B. E., C. S. KEENER AND B. A. FORD. 1997. Anemone. Pp. 139-158 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, Vol. 3. Oxford University Press, New York, NY. KEARNEY, T. H. AND R. H. PEEBLES. 1951. Arizona flora. University of California Press, Berkeley, CA. Munz, P. A 1974. A flora of southern California. University of California Press, Berkeley, CA. SAN DIEGO NATURAL HISTORY MUSEUM, 2011. The flora of Baja California. San Diego Natural History Museum, San Diego, CA. Website http:// bajaflora.org/ [accessed 3 March 2011]. WIGGINS, I. 1980. The flora of Baja California. Stanford University Press, Stanford, CA. WILKEN, D. H. 1993. Anemone. Pp. 912-913 in J. C. Hickman (ed.), The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. 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