Historic, Archive Document Do not assume content reflects current scientific knowledge, policies, or practices. 750 NP o® VOLUME 49, NUMBER 1 JANUARY-MARCH 2002 EFFECTS OF FIRE ON THE REPRODUCTIVE BIOLOGY OF AGAVE PALMERI (AGAVACEAE) [ELE SSS OTIS DB resco SB Ee PE EE SI Re 1 ADULT SEX RATIO OF ARCEUTHOBIUM GILLII (VISCACEAE) Robert L. Mathiasen and Carolyn M. Daugherty .............cccccccceeeeeeeveneeees 12 ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE (POLYGONACEAE), A NEW VARIETY FROM THE SOUTHERN SIERRA NEVADA, CALIFORNIA | ROSE OING NOTE OO oN I RR eR TI Te a 16 HESPEROYUCCA WHIPPLEI AND YUCCA WHIPPLEI (AGAVACEAE) Jeffrey A. Greenhouse and JON, StrOMOCB ecceccccccess-c00ceese0s.04000000nenaneees- 20 SYMPATRY BETWEEN DESERT MALLOw, EREMALCHE EXILIS, AND KERN MALLow, E. KERNENSIS (MALVACEAE): MOLECULAR AND MORPHOLOGICAL PERSPECTIVES Katarina Andreasen, Ellen A. Cypher and Bruce G. Baldwin................. 22 POLLINATION OF CYTISUS SCOPARIUS (FABACEAE) AND GENISTA MONSPESSULANA (FABACEAE), Two INVASIVE SHRUBS IN CALIFORNIA Ingrid M. Parker, Alexandra Engel, Karen A. Haubensak and Karen Goodell ....... ARE A PTS Bi os en scwecececcsecee DS FOxTAIL PINE IMPORTANCE AND CONIFER DIVERSITY IN THE KLAMATH MOUNTAINS AND SOUTHERN SIERRA NEVADA, CALIFORNIA Andrew J-Eekert and JOGA sO: SAWCT ie FB Ss ISP ne ono 00 ET wae 33 NOTEWORTHY BRYOPHYTE RECORDS FROM THE MOJAVE DESERT Lloyd R. Stark, Alan T. Whittemore and Brent D. Mishler................00000+ 49 COPNTETEGRNA, oo ss ccs Ee re cn scene ceccecettocss 54 © TERN yea ses I sat soc oi Ue hc oa wadevounaneedcbboveSes 54 IB YATS1 0) Sornse acer On cl ere ome mi 7A 220: SEEN ARR nEet12)\ 54 AN TONGTONINVN es eso ce oe ook Ne Og ee ee ed ey ee ccateeu sche epee 55 INTER RY DED ICC) ck yang tee ee Sen err eg 1061) 0 Ae Oe an 54 CO) EE Oe en ate inc Sn BS NRE SM Fld Sec anuys aia uc tica seuunb suneentceuies 58 THE MANZANITAS OF CALIFORNIA, ALSO OF MEXICO AND THE WORLD, BY Putte V. WELLS MuachaelC. Vasey Gnd WV ThOMGS POLK «.s50502.03ssccsssesesnecssssancteeestespunsiee 46 ILLUSTRATED FIELD GUIDE TO SELECTED RARE PLANTS OF NORTHERN CALIFORNIA, EDITED BY GARY NAKAMUA AND JULIE KIERSTEAND NELSON TEATRO) eS CNN Cre 5 ee ee arate Des SEN CR co Sistah AS caidas iedtsis donut as Wee Uwaus 48 Monocots III/Grasses IV .. RECEIVED BY: re ES fo ee 15 FERRVATIUINI elo oeSiceressiccdeonton. estan: leartsias a 59 INDEXING (2 4) PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY Maprono (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. PostMAsTER: Send address changes to MApRONO, Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. Editor—Dr. JOHN CALLAWAY Dept. of Environmental Science University of San Francisco 2130 Fulton Street San Francisco, CA 94117-1080 callaway @usfca.edu Book Editor—Jon E. KEELEY Noteworthy Collections Editors—DieTER WILKEN, MARGRIET WETHERWAX Board of Editors Class of: 2002—NorMAN ELLSTRAND, University of California, Riverside, CA Cara M. D’ Antonio, University of California, Berkeley, CA 2003——-FREDERICK ZECHMAN, California State University, Fresno, CA Jon E. Kee.ey, U.S. Geological Service, Biological Resources Division, Three Rivers, CA 2004—Davip M. Woon, California State University, Chico, CA INGRID PARKER, University of California, Santa Cruz, CA 2005—J. Mark Porter, Rancho Santa Ana Botanic Garden, Claremont, CA Jon P. REBMAN, San Diego Natural History Museum, San Diego, CA CALIFORNIA BOTANICAL SOCIETY, INC. OFFICERS FOR 2001—2002 President: Bruce BALDwin, Jepson Herbarium and Dept. of Integrative Biology, 1001 Valley Life Sciences Bldg. #2465, University of California, Berkeley, CA 94720. First Vice President: Rop Myatt, San José State University, Dept. of Biol. Sciences, One Washington Square, San José, CA 95192. rmyatt @email.sjsu.edu Second Vice President: PETER Fritscu, Dept. of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. pfritsch @calacademy.org Recording Secretary: DEAN KELCcu, Jepson and University Herbarium, University of California, Berkeley, CA 94720. dkelch @sscl.berkeley.edu Corresponding Secretary: SUSAN BAINBRIDGE, Jepson Herbarium, 1001 VLSB #2465, University of California, Berkeley, CA 94720-2465. 510/643-7008. suebain@SSCL.berkeley.edu Treasurer: Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. The Council of the California Botanical Society comprises the officers listed above plus the immediate Past President, R. Joun Litre, Sycamore Environmental Consultants, 6355 Riverside Blvd., Suite C, Sacramento, CA 95831; the Editor of Maprono; three elected Council Members: BiAN Tan, Strybing Arboretum, Golden Gate Park, San Fran- cisco, CA 94122; James SHEvock, National Park Service, 1111 Jackson St., Suite 700, Oakland, CA 94607-4807. 510/ 817-1231; ANNE BRADLEY, USDA Forest Service, Pacific Southwest Region, 1323 Club Drive, Vallejo, CA 94592. abradley @fs.fed.us; Graduate Student Representative: KirstEN M. FisHer, Jepson Herbarium, University of California, Berkeley, CA 94720. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). MADRONO, Vol. 49, No. 1, pp. 1-11, 2002 EFFECTS OF FIRE ON THE REPRODUCTIVE BIOLOGY OF AGAVE PALMERI (AGAVACEAEB) Liz A. SLAUSON Scottsdale Community College, 9000 E. Chaparral Rd., Scottsdale, AZ 85256-2626 ABSTRACT Fire is an important management tool that is used increasingly to restore natural composition, structure, and processes in semi-arid grasslands, woodlands, and forests of the southwestern U.S. I investigated the effects of fire on floral resources, fruit and seed set, and survivorship of Agave palmeri, an important food source of the endangered lesser long-nosed bat. Nectar production and sugar concentration, pollen and nectar standing crops, and fruit and seed production were measured in flowering plants with leaves damaged by fire and compared with unburned plants at three sites in two different human-ignited fires. In general, no significant differences were found in total nectar production, nectar sugar concentration, standing pollen crops, or fruit and seed set between burned and unburned plants. Standing nectar crops were slightly smaller than total nectar production amounts, but large amounts of nectar and pollen re- mained available at dawn in both burned and unburned plants. Initial mortality measured across all size classes at one site was only 3.3%. Although levels of burn damage relative to plant size were quite variable, plants with greater damage (61—100%) tended to be <0.6 m in height and diameter. These results indicate that fire did not appreciably decrease food resources of the lesser long-nosed bat or the repro- ductive resources and survivorship of A. palmeri. The rocky, low fuel habitats preferred by A. palmeri combined with certain morphological and physiological adaptations of the plant, such as a rosette shape and storage of accumulated carbohydrates in the center of the rosette, appear to protect the majority of stored resources within the plant’s center during fire events. Key words: Agave palmeri, fire, Leptonycteris curasoae yurbabuenae, lesser long-nosed bat. INTRODUCTION Agave palmeri Engelm. is a perennial, rosette- shaped leaf succulent, and is widespread in desert scrub, oak savanna, and oak woodland communities of the Southwest Borderlands: the international four-corners area of Arizona, New Mexico, Sonora, and Chihuahua. Prescribed fire is one management tool that is being tested to control the conversion of grasslands into shrubland communities, to re- duce high fuel loads in woodland communities, and to return communities of the Southwestern Border- lands to pre-settlement states (Allen 1996; Edmins- ter 1996). Although the pollen and nectar of A. pal- meri are major food sources of the endangered less- er long-nosed bat (Leptonycteris curasoae yurba- buenae) (Hayward and Cockrum 1971; Howell 1972; Howell and Roth 1981; Slauson 2000), little is known about the effects of fire on A. palmeri, and in particular, on its production of food resourc- es for floral visitors. Agave palmeri has a prolonged juvenile period that may last 20—40 years, during which time water and carbohydrates accumulate in the leaves. Once maturity is reached, plants are monocarpic, flow- ering only once and then dying. Mature plants range in size from 0.5—1.6 m tall and 0.5—2.5 m wide with 56-124 leaves (Noble and Smith 1983; Slauson 2000). The inflorescence, or flowering Stalk, is a large, open panicle that varies from 2-8 m in height with 10—30 umbels (flower cluster on side branch of inflorescence) (Slauson 1996; Hodg- son 1999), and produces on average 1500-2200 flowers over the flowering season (Slauson 2000). Anthers are sizeable (8.7—20 mm in length) and produce large amounts of pollen if not removed on the night of dehiscence by bats, moths, or rain; pol- len is usually harvested the following morning by various bees (Slauson 2000). Flowers secrete nectar nocturnally over a 6-day flowering period and pro- duce an average of 2.5 ml of total nectar, with peak production on the second day (Slauson 1999, 2000). These large quantities of pollen and nectar attract many animals (Slauson 2000), including the lesser long-nosed bat. Nocturnal visitors have the ‘‘first crack’? at pollen and newly secreted nectar. The lesser long-nosed bat was federally listed as endangered in 1988 due to apparent low and de- clining numbers, disturbance of roosts, and an in- terdependence with its food resources (Shull 1988), but its status as endangered is controversial (see Cockrum and Petryszyn 1991). Several species of columnar cacti and agaves provide a “‘nectar cor- ridor’’ for the lesser long-nosed bats, from spring as they migrate north from Central America and Mexico, through fall when they return to southern roosts (Gentry 1982; Fleming et al. 1993). In late July and early August, substantial numbers of lesser long-nosed bats migrate to higher elevations in southeastern Arizona (Cockrum 1991) where their primary food source from July through September is A. palmeri (Howell 1972; Howell and Roth 1981). In describing this mutualistic relationship, p) MADRONO Howell and Roth (1981) suggested that A. palmeri was also “‘strongly dependent’? upon the lesser long-nosed bat for pollination. More recently, Slau- son (2000) has shown that this mutualistic relation- ship is asymmetrical; bats depend on this agave species part of the year for food, but A. palmeri does not require bats for adequate sexual reproduc- tion. Episodic fires were common throughout most ecosystems of the Southwest Borderlands for at least 300—400 years prior to ca. 1900 (Baisan and Swetnam 1990; Swetnam and Baisan 1996a, b), in- cluding those occupied by A. palmeri. Kaib (1998) has shown desert grasslands burned approximately every 5—10 years, and ranged between 10 to hun- dreds of km? in size, while canyon pine-oak forests burned every 5—9 years and covered areas of at least 50 km’. A drastic reduction of these wide- spread, episodic surface fires occurred north of the border between ca. 1870—1900, initially as a result of intensive grazing that decreased fuels and the ability of fire to spread across large landscapes (Bahre 1991; McPherson et al. 1993; Allen 1996; Swetnam and Baisin 1996a, b). Continued grazing, land use, and fire suppression practices by govern- ment agencies throughout the 20th century resulted in the virtual extinction of natural disturbance fires in southwest ecosystems (Bahre 1991; Allen 1996; Swetnam and Baisan 1996a). These practices have contributed to many changes in both the structure and function of these communities, ranging from shrub invasion of desert grasslands due to a lack of fine fuels to carry low intensity fires (Hastings and Turner 1965; Humphrey 1987; McPherson 1995), to severe risk of catastrophic and stand-replacing wildfires in forests due to great fuel accumulations (Cooper 1960; Covington and Moore 1994). Many succulents tolerate fire to some degree (Thomas and Goodson 1992), but desert grassland fires have also been reported to kill succulent plants (Niering and Lowe 1984; Nobel 1988), particularly the smaller size classes. McLaughlin and Bowers (1982) reported that large succulent plants that ini- tially survived fires had increased mortality in later years. Several other factors besides plant size may affect the responses of succulents to fire including the size and patchiness of the fire, the time of year fire occurs, fuel conditions, weather conditions, to- pography, and plant morphology and physiology (McPherson 1995). Little data are available regard- ing the effects of fire on agaves. In simulated grass- land fire experiments, A. gigantensis had 0% mor- tality after 14 months with fuel densities up to 1600 g m ~~? and temperatures of 400—600°C (these tem- peratures are normally expected in intense grass- land fires) (Thomas and Goodson 1992). Although the leaf tips collapsed at fuel densities of 400 g m ~~ and damage increased as fire intensity increased, the center of the rosettes remained unburned. In field surveys of semi-desert grassland sites in southern Arizona that had burned in wildfires in the [Vol. 49 prior 18 months, mortality from fire was 18% in A. palmeri (Thomas and Goodson 1992). Of the re- maining living plants, 83% exhibited regrowth from the apical meristem, whereas 17% survived un- burned in refugia. These refugia were created by either the patchy nature of the fire, which skipped Over areas with adequate fuels, or by rocky areas with little flammable material. The prolonged juvenile period of agaves results in large amounts of stored resources that are used for flower (including nectar and pollen), fruit, and seed production. Once flowering is initiated, a large and irreversible translocation of stored resources occurs from the rosette to the developing inflores- cence. Death results presumably because resources normally reserved for growth and maintenance are mostly allocated to reproduction. Nobel (1977) ob- served that A. deserti Engelm. diverted over 68% of its stored biomass to a developing inflorescence. No data are available regarding how the loss of stored resources from fire damage in reproductive agaves affects nectar and pollen production or sub- sequent fruit and seed set. Research questions. 1 studied aspects of the floral biology of Agave palmeri relevant to nectar bat vis- itation immediately following two summer fires. These fires, one prescribed and one accidental, burned patchily and caused little immediate mor- tality of mature agave plants. Plants that were re- productive the year of each fire had partially or ful- ly emerged inflorescences at the time of the fires, and either were flowering or flowered soon after. I compared reproductive burned plants with un- burned plants and asked (1) did nectar production rate or concentration differ between the two treat- ment classes? (2) What was the standing crop of nectar and pollen at dawn, after possible bat visi- tation? (If floral rewards of populations were close- ly cropped by bats, then an adverse effect of burn- ing on nectar production would be important. A surplus in both nectar production and standing crops would indicate that floral foods were not in limited supply for nocturnal visitors. Differences between burned and unburned plants could indicate production differences or discrimination by noctur- nal visitors.) (3) Did fruit and seed set differ be- tween burned and unburned plants? (A significant decrease in fruit or seed set could have considerable impacts on future floral resources for bats.) (4) Was burn damage of plants related to their size, specif- ically the height and diameter of the rosette? METHODS Study sites. Fieldwork was conducted at two dif- ferent fire sites, a prescribed burn and an accidental human-ignited fire. The prescribed burn (known as the Maverick Burn) was conducted on June 24—25, 1997 by the U.S. Forest Service on the southern edge of the Peloncillo Mountains (Fig. 1). Approx- imately 8000 acres burned in a mosaic pattern with- 2002] Arizona New Mexico Peloncillo Mtns. Animas Mts. Chihuahua Sonora Fic. 1. Locations of CE GT (Maverick Burn), and MF (Gray Ranch) study sites. in the 17,000 acres designated as the primary burn area (Encinas 1997). Two study sites were chosen within the primary burn area that represented typ- ical habitats for A. palmeri (Gentry 1982). The first study site was located in the southern portion of the burn area on Cowboy Flats (CF site, lat 31°26'N, long 109°2'W, elevation ~1585 m) near a small lesser long-nosed bat roost. This site was on a rocky, south-southwestern facing hillside and mesa top in an oak savanna community. Plant species composition included Quercus emoryi, Q. oblon- gifolia, Juniperus monosperma, Prosopis velutina, Yucca schotti, Nolina microcarpa, Dasylirion wheeleri, Fouquieria splendens, Calliandra sp., Gutierrezia sarothrae, Bouteloua gracilis, B. cur- tipendula, B. hirsuta, B. radicosa, Hilaria belan- geri, Erogrostis intermedia, Muhlenbergia sp., Ly- curus phleiodes, Aristida spp., Schizachrium cir- ratum, and Heteropogon contortus. The second study area was located along the northern border of the burn area along Geronimo Trail (GT site, lat 31°32’N, long 109°2'W, elevation ~1675 m), and was characterized by flat to steep (0O—45°) slopes with south-southwestern exposures and rocky soils in an oak savanna/oak woodland community. Plant species included Q. emoryi, Q. oblongifolia, J. monosperma, J. deppeana, P. velutina, Arctostaph- ylos pungens, Y. schotti, G. sarothrae, B. gracilis, B. curtipendula, B. hirsuta, B. radicosa, H. belan- geri, E. intermedia, Aristida spp., S. cirratum and M. emersleyi. The accidental human-ignited fire started in Mexico in May 1999, crossed the international bor- der, and burned approximately 22,000 acres in a mosaic pattern over a 2-day period on the Gray SLAUSON: FIRE EFFECTS ON AGAVE PALMERI 3 Ranch in southeastern New Mexico (Fig. 1). The third study site was located on the Gray Ranch near McKinney Flats (MF site, lat 31°23’N, long 108°42’'W, elevation ~1570 m) on a south-facing, rocky slope in a semi-arid grassland/oak savanna community. Plant species included Q. emoryi, Q. oblongifolia, P. velutina, N. microcarpa, Juniperus sp., Cylindropuntia spinosior, G. sarothrae, B. gracilis, B. curtipendula, B. hirsuta, B. radicosa, H. belangeri, Aristida spp., and S. cirratum. Agave populations (all size classes excluding seedlings) at study sites ranged from approximately 1000—3000 plants per ha. Although several plots were identified as poten- tial study sites prior to the Maverick fire, ignitions were initiated across broad landscape areas rather than igniting individual study plots, and none of the potential study plots burned. Therefore, thorough pre-treatment characterization of plots was not pos- sible. Due to the large areas of steep and inacces- sible terrain, the mosaic pattern of the fire, the patchy distribution of agaves, and limitations of sampling nectar and pollen with 12-foot ladders, only two suitable sites that actually burned could be located within the primary burn area. At these sites, fire behavior was characterized by partial to complete understory consumption (Clark 2000). The Gray Ranch fire was accidental, and only one accessible agave population burned. Research was conducted at both fire sites during late July and August during the peak flowering pe- riod of A. palmeri and when migrating lesser long- nosed bats are normally present in southeastern Ar- izona. Work was conducted August 4—9, 1997 at the CF site, August 11—15, 1997 at the GT site, and July 20—22 and August 2—4, 1999 at the MF site. Nectar and pollen studies. 1 studied nocturnal nectar production in bagged, or exclosed, flowers to determine whether this important floral reward was affected by fire. I also measured standing crops of nectar and pollen at dawn in unbagged, or open, flowers. Standing crops reflect both the effects of harvesting by animals and production of floral re- wards by flowers. Plant selection was limited to plants whose flowers could be reached with 12-foot ladders in burned and unburned areas at each site. A plant was classified as a “‘burned plant” if >1% of the rosette had tissue death due to fire. Rosette height, width, and number of umbels were mea- sured for each study plant. For study plants in burned areas, burn damage (% of rosette damaged by fire) was scored from 0 to 6: 0 = no burn dam- age, 1 = 1-20%, 2 = 21-40%, 3 = 41-60%, 4 = 61-80%, 5 = 81-—100%, and 6 = entirely dead. Sample sizes for nectar and pollen studies at all sites were restricted due to the mosaic pattern of the fire, the patchy distribution of agaves on pri- marily inaccessible and rocky terrain, and the num- ber of plants in flower that could be reached with 12-foot ladders. In general, all plants that could be 4 MADRONO reached at a site were sampled. Total nectar pro- duction, nectar sugar percentage, and standing nec- tar and pollen crops were measured on six plants each in burned and unburned plots at the CF site and nine plants at the unburned GT site. Flowers on most plants in the burned GT plot were too high to reach with ladders, and only 2—3 plants could be sampled, depending on the day and appropriate flower stage availability. At the MF site, 20 and 10 plants were measured in the burned and unburned plots, respectively. To decrease any positional ef- fects of nectar volume, nectar sugar concentration or standing crops, experimental flowers were locat- ed on umbels positioned in the middle section of the inflorescence, whenever possible. For studies of total nectar production and nectar sugar concentration at the CF and GT sites, 5-10 predehiscent (day 1 of 6-day flowering period) flowers on one umbel were numbered, enclosed with a nylon mesh bag prior to dusk, and nectar was allowed to accumulate until dawn (nectar is produced only during the night). Anthers of pre- dehiscent flowers generally split open and dehisced pollen shortly after dusk, entering the dehiscent (day 2) floral stage. Dehiscent stage flowers were used in nectar and pollen experiments as nectar pro- duction is greatest in dehiscent flowers, and pollen is only available during the dehiscent stage (Slau- son 2000). Plants were sampled for three successive days for total nectar production studies (different flowers sampled each day) and one day for nectar sugar concentration studies. Nectar production was measured by withdrawing the nectar present in the floral tube at dawn with a tuberculin syringe and blunt end needle. Nectar sugar concentration (per- cent sucrose equivalents on a weight/weight basis) was measured in the field at dawn with a hand-held refractometer. Data collection methods for total nectar production and nectar sugar concentration were modified for the MF site in an attempt to in- crease the number of plants sampled for compari- son to standing nectar crops (see below). Plants were sampled one day only and data collection was spread over a 3-day period (4—7 plants per treat- ment were sampled each morning at dawn). To de- termine if nectar resources decreased significantly over time, total nectar production and nectar sugar concentration measurements were repeated on the same study plants two weeks later (August 2—4, 1999). Due to the difference in sampling method- ology, fire sites were analyzed separately. To indirectly determine the degree of nectar use by nocturnal floral visitors (bats and moths), stand- ing nectar crops (amount of nectar present at dawn in flowers left available to visitors) were measured and compared to total nectar production (exclosed flowers) at all three study sites. In order to most accurately measure dawn standing crops, sampling was begun as close to dawn as possible (0.54 ml) were available at dawn at all sites. A significant interaction was present in nectar production be- tween the burn treatment and standing nectar crop vs. total nectar production at the CF site (one-way repeated measures ANOVA, F,,) = 5.765, P = 0.037), and as nectar production was averaged across the burn treatment, it was not advisable to interpret the standing crop vs. total nectar produc- tion test. Data were subsequently analyzed by run- ning paired t-tests on standing nectar crop and total nectar production data by burn treatment. Standing nectar crop (mean + SE = 0.565 + 0.033 ml) was significantly lower than total nectar production (mean + SE = 0.662 + 0.027 ml) in burned plants (t-test, ¢ = —1.767, P = 0.017, alpha/2 (0.025) to maintain Type I error), but no significant difference was found in nectar production between standing nectar crop (mean + SE = 0.546 + 0.029 ml) and total nectar production (mean + SE = 0.558 = 0.028 ml) in unburned plants (t-test, t = —0.244, P = 0.646, alpha/2 (0.025) to maintain Type I error). At the GT site, no significant differences in nectar < Fic. 2. Mean total nectar production (ml) of dehiscent A. palmeri flowers in burned and unburned plots, (a) CF site, (b) GT site, and (c) MF site. Vertical lines = 1 SE. 2002] production were found between standing nectar crop (mean + SE = 0.612 + 0.031 ml) and total nectar production (mean + SE = 0.711 = 0.018 ml) in unburned plants (one-way repeated measures ANOVA, F,, = 1.101, P = 0.371) or in nectar production between days (one-way repeated mea- sures ANOVA, F,, = 3.927, P = 0.081). Data for standing nectar crop and total nectar production in burned plants were not analyzed due to low sample size (n = 2) and missing paired data values (flowers were not available for both open and exclosed nec- tar production on all three nights for the three plants with flowers that could be reached). At the MEF site, no significant differences in nectar production were found between burned and unburned plants (one- way repeated measures ANOVA, F)5, = 2.764, P = (0.108) or standing nectar crop and total nectar production (one-way repeated measures ANOVA, Fi53 = 0.791, P = 0.381) (standing crop mean + SE = 0.515 + 0.038 ml, total nectar production mean + SE = 0.557 + 0.042 ml). Nectar sugar concentrations of standing nectar crops ranged from 16.7—29.6% at the CF site and 16.2—21.9% at the GT site. Nectar sugar concentra- tions were significantly higher at the CF site (Proc Mixed procedure, F,5, = 7.11, P = 0.015) and in standing crop flowers at both sites (Proc Mixed pro- cedure, F',, = 8.70, P = 0.021) (CF site: standing crop mean + SE = 21.8 + 1.1%, total nectar pro- duction mean + SE = 20.6 + 0.9%; GT site: stand- ing crop mean + SE = 19.2 + 0.6%, total nectar production mean + SE = 17.5 + 0.4%). At the MF site, standing nectar sugar concentrations ranged from 12.6-19.5%. No significant differences in nectar sugar concentrations were found between burned (mean + SE = 15.8 + 0.3%) and unburned plants (15.0 + 0.5%) (one-way repeated measures ANOVA, F453, = 2.220, P = 0.147) or between standing nectar crop (mean + SE = 15.6 + 0.3%) and total nectar production (mean + SE = 15.5 + 0.2%) (one-way repeated measures ANOVA, F, 5. = 1.167, P = 0.289). Fruit and seed set. Burned and unburned plants had similar levels of fruit (17-22%) and seed set (19-23%) (Fig. 3). No significant differences were found in fruit set between study sites (one-way AN- OVA, Fy., = 1.915, P = 0.154). Fruit set was somewhat lower in burned plots, but not signifi- cantly (one-way ANOVA, F,,, = 0.294, P = 0.589). Seed set was slightly higher in two of the three burned plots, but differences were not signif- icant (one-way ANOVA, F\ 45 = 0.641, P = 0.427). Seed set at the CF site was significantly lower than the GT and MEF sites (one-way ANOVA, Bonfer- roni test, P < 0.05). Mortality and demography. Overall mortality of plants in demography plots measured two months after the Gray Ranch fire (MF site) was 3.3% (Plot 1 = 4.6%, n = 194, Plot 2 = 2.5%, n = 355, Plot 3 = 3.9%, n = 276, Plot 4 = 3.0%, n = 459). Mean SLAUSON: FIRE EFFECTS ON AGAVE PALMERI 7 50 45 Population 40 & CF 35 &@ GT O MF 30 25 20 15 10 Fruit set (%) burned unburned Treatment a- 4 | burned unburned Treatment Seed set (%) AAS (eo) Fic. 3. Mean fruit and seed set (%) of A. palmeri in burned and unburned plots, CE GT, and MEF sites. Vertical lines = 1 SE. burn damage + SE was 3.1 + 0.03 (41-60% of rosette damaged) (range of plots = 3.4 + 0.06 to 2.7 + 0.05) with approximately 37% of plants fall- ing into this damage class (Fig. 4). Burn damage classes 4 (22% of plants) and 2 (16% of plants) were the next most frequent damage classes. A multinomial logit model (Agresti 1990) relating burn class damage to rosette height and width cor- rectly predicted damage class for only 25% of ob- servations. The variability in this relationship is shown in Figure 5. Although median plant size across all levels of burn damage did not differ greatly, the range was quite variable. Rosette height and width values associated with burn damage clas- ses 23 were more concentrated than those of burn damage classes 1-2. Rosette height and width size class distributions illustrate the relatively young age of the population (Fig. 6). Greater than 90% of the population was <0.3 m in height and width. 8 MADRONO [Vol. 49 600 1000 900 0.4 0.7 500 800 0.6 2 a 20 ou = 400 age & 05% row = co 600 2 ° 300 S ° 500 0.4 8 ( z= 300 o> 01 = Ons 100 aut 100 0.1 0 0 0.0 0 1 20 40 60 80 dead 0.0 0.1 02 03 04 05 06 0.7 08 09 1.0 Burn damage (%) Fic. 4. Burn damage (% of rosette damaged by fire) of A. palmeri at the MEF site. 0 = no burn damage, 1 = 1-20%, 2 = 21-40%, 3 = 41-60%, 4 = 61-80%, 5 = 81—100%, 6 = dead. DISCUSSION Burning did not appear to negatively impact the production of nectar in A. palmeri. Mean total nec- tar production for all sites and times sampled (range + SE = 0.46—-0.71 + 0.5 ml) was similar to or greater than previous reports of nectar production Rosette height Rosette width Burn damage 0 0.5 1.0 1.5 Rosette height and width (m) Fic. 5. Relationship between plant size (m) and burn damage (% of rosette damage by fire) of A. palmeri, MF site. 0 = no burn damage, 1 = 1—20%, 2 = 21-40%, 3 = 41-60%, 4 = 61-80%, 5 = 81-100%, 6 = dead. Lower box plot of each pair indicates rosette height, upper box plot indicates rosette width. Length of box plot = range within 50% of values fall, center notch = median, box edges = first and third quartiles, whiskers = values within the first quartile — (1.5 X midrange) and the third quartile + (1.5 X midrange), asterisks = values between first quar- tile — (1.5 to 3.0 X midrange) and third quartile + (1.5 to 3.0 X midrange), and open circles = values third quartile + (3.0 X midrange). Rosette height (m) Number of plants Jeg Jed uoimodold OH) 0:1 0 OL OF Oks OLS (0.7 O13 O.2 710 Rosette width (m) Fic. 6. Size class distribution of A. palmeri, MF site. n = 1298 plants. in dehiscent flowers of unburned plants (Howell 1979; Slauson 1999, 2000). The significantly lower nectar production of burned plants at the GT site may be a result of sampling error due to the low sample size (n = 3 plants). Nectar sugar concentra- tions were not significantly different between burned and unburned plants at any site, and values were well within the range reported in previous studies of unburned plants (Howell 1979; Slauson 1999, 2000). Nectar production and nectar sugar concentrations decreased over time in both burned and unburned plants, and this trend suggests that nectar resources may naturally decrease during the flowering period as stored resources are depleted and fruit and seed production increase. Pollen pro- duction did not appear to be affected adversely as large amounts of pollen were present on anthers at dawn at all sites and sampling times. Standing nectar crops at dawn were smaller than total nectar production in all treatments at all sites 2002] (only significantly lower at the burned CF site), pre- sumably as a result of nectar use by moths and bats. However, at all study sites large amounts of nectar (>0.54 ml) and pollen (score = 5) were available in standing crop flowers at dawn, and indicate that food resource availability was not a limiting factor for either nocturnal (bats or moths) or diurnal vis- itors at any study sites during the times sampled. Standing crop results were similar to previous re- ports from other populations (Slauson 2000). The study sites examined did not appear to con- tain the fuel loads required to support a fire of the intensity needed to kill mature (reproductive) aga- ves or cause significant initial mortality when all size classes were considered. Although the majority of the population was quite small in size at the MF site (Fig. 6), mortality was <4%. Mean burn dam- age of mature plants was moderate at all sites (ap- proximately 10—43% of rosette burned), and in no case did mean damage exceed class 3 (41-60% of rosette burned). Relatively few large, non-repro- ductive plants (>0.6 m in height or diameter) had burn damage >60% (Fig. 5), and mean burn dam- age of all size classes at the CF site was approxi- mately 43%, despite the fact that over 90% of the population was <0.3 m in height and diameter. The low to moderate burn damage and high ini- tial survivorship of agaves may be due in part to their preferred habitats. Agave populations of mod- erate to high densities are characteristically found in xeric sites with rocky soil surfaces where com- petition with other plants is low (Gentry 1982). These rocky, low fuel habitats may serve as a re- fugia from fire for some agaves, especially smaller agaves which may be protected near the bases of surface rocks and cobbles. Plants located in less rocky, denser grassland or woodland habitats with increased fuel loads would presumably sustain greater damage. The greater burn damage observed in plants at the GT site (41-60%) may have been due to larger amounts of available fuels present in the surrounding oak savanna and oak woodland community. Robinett and Barker (1996) noted that frequent seedling establishment of A. palmeri oc- curs around dried, dead adult plants, and fuel loads created by dead adults can produce intense heat in a fire that kills surrounding seedlings. The vari- ability in the fuel loads of individual agave micro- habitats most likely explains the unpredictability of burn damage as a function of plant size observed in this study, especially in the smaller size classes (Fig. 5). Certain morphological and physiological adap- tations may also lessen the effect of fire on agaves. A number of agave species may resprout from rhi- zomes after fire (Gentry 1972), although most pop- ulations of A. palmeri are not rhizomatous (Slauson personal observation). Agaves may also benefit from the release of nutrients after a fire due to their Shallow root system (Gentry 1982). The rosette form of agaves to some degree protects the apical SLAUSON: FIRE EFFECTS ON AGAVE PALMERI 9 meristem and the majority of stored resources lo- cated in the center of the plant. Reproductive aga- ves may have an even greater advantage: beginning in late winter and early spring, stored water and carbohydrates in reproductive agaves are translo- cated from the leaves to the center of the rosette where they are utilized for the development of the inflorescence (Nobel 1977; Tissue and Nobel 1990). Concentrated in the center of the plant, the accumulated resources have maximum protection from fire, while the outer leaves, which are most susceptible to fire damage, can burn with little overall loss of stored carbohydrates. The timing of inflorescence emergence may also be important in reducing fire damage in reproductive agaves. Most natural lightning strikes that result in fires occur from May—August (Sellers and Hill 1974; Swetnam and Baisan 1996b). By this time, flower stalks are generally taller than the surrounding vegetation in grassland communities and above the fire zone (Slauson, personal observation). Spring fires or fires in denser oak woodland communities where the fire zone reaches into tree canopies could po- tentially damage emerging inflorescences more se- verely. Low fruit set is common in outcrossing, her- maphroditic plants such as agave (Sutherland and Delph 1984). Sutherland (1982) found that mean fruit set per inflorescence in paniculate agaves is consistently around 20% despite hand or open pol- lination treatments, suggesting that fruit set is pri- marily resource limited. Slauson (2000) studied fruit and seed set in three different open-pollinated populations of A. palmeri in Arizona, and also found fruit set to average around 20% while seed set varied from 26—33%. Fire did not appear to af- fect reproductive output in this study as no signif- icant differences in fruit and seed set were found between burned and unburned plants, and results (Fig. 3) were very similar to previous reports of unburned plants. The rosette shape and transloca- tion of resources in reproductive agaves appear to be important adaptations to fire that protect stored resources critical for reproduction. The significantly lower seed set at the CF site was observed in both burned and unburned plants, and suggests the cause is most likely a result of pollinator and/or resource availability at the site and not a result of burn treat- ment. Periodic fires may be important in promoting germination and establishment events of A. palmeri by reducing competition and opening up germina- tion sites, especially in dense grassland/woodland habitats. Germination of A. palmeri generally oc- curs in mid-late summer after monsoon rains have begun. During July 1999 (two months after the fire) approximately four inches of precipitation fell at the ME site, and during establishment of demog- raphy plots in burned areas in late July—early Au- gust, 16 seedlings were observed. These plants were assumed to be seedlings as they were <40 X 10 MADRONO 20 mm in height and diameter, and had no burn damage despite being located near burned plants. A cursory search for seedlings was conducted in several unburned areas adjacent to the burned site, but no seedlings were found. CONCLUSIONS Prescribed natural and human-ignited fires are important ecosystem management tools used to al- ter community composition and forage conditions, decrease fuel accumulation, and reduce the poten- tial of catastrophic, stand-replacing wildfires. Al- though historic changes in the southwestern U.S. landscape have been significant due to fire exclu- sion, the return of fire in the sites studied did not appreciably impact nectar and pollen production, fruit and seed set, or initial mortality of A. palmeri. Standing crop results indicated that at least through mid-August in the sites examined, food resources of lesser long-nosed bats were not limited as a re- sult of fire. Habitat preferences and several mor- phological and physiological adaptations appear to adequately protect the majority of stored resources of A. palmeri during fire events, although fuel loads within individual microhabitats of agaves can result in variable fire damage to rosettes. Further study is needed to more clearly understand how fire affects population dynamics and mortality of A. palmeri beyond the initial post-fire period. ACKNOWLEDGEMENTS The author thanks J. McAuliffe, P. Scott, C. Edminster, and two anonymous reviewers for helpful guidance and comments on the manuscript; J. Borgmeyer, M. Johnson, K. Mueller, K. Rice, S. Ahearn, D. Hansen, N. Grant, S. Garrison, T. Omar, K. Smith, and volunteers and staff of the Desert Botanical Garden for field and laboratory as- sistance; P. Sundt for assistance with plant species data, R. King, Rocky Mountain Research Station, for statistical advice and consultation; B. McDonald of the Sycamore Ranch and Ben Brown and the Animas Foundation of the Gray Ranch for logistical and study site location assis- tance; and Ginny and Dave Dalton for assistance in bat identification and observations. This research was sup- ported in part by funds provided by the Rocky Mountain Research Station, U.S. Department of Agriculture, Forest Service, and the Desert Botanical Garden. LITERATURE CITED AGRESTI, A. 1990. Categorical data analysis. John Wiley & Sons, New York. ALLEN, L. S. 1996. Ecological role of fire in the Madrean Province. 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Historical fire regime patterns in the Southwestern United States since AD 1700. Pp. 11—32 in C. D. Allen (technical editor), Fire effects in southwestern forests, proceedings of the 2nd La Mesa Fire Symposium. General Technical Re- port-RM-GTR-286. U.S.D.A. Forest Service, Rocky Mountain Forest and Range Experiment Station, Ft. Collins, CO. THOMAS, P. A. AND P. GoopDson. 1992. Conservation of succulents in desert grasslands managed by fire. Bi- ological Conservation 60:91—100. TISSUE, D. T. AND P. S. NoBEL. 1990. Carbon relations of flowering in a semelparous clonal desert perennial. Ecology 71:273-281. MADRONO, Vol. 49, No. 1, pp. 12—15, 2002 ADULT SEX RATIO OF ARCEUTHOBIUM GILLIT (VISCACEAE) ROBERT L. MATHIASEN School of Forestry, Northern Arizona University, Flagstaff, AZ 86011 CAROLYN M. DAUGHERTY Geography and Public Planning, Northern Arizona University, Flagstaff, AZ 86011 ABSTRACT The adult sex ratio of Arceuthobium gillii was determined from five populations in southern Arizona and one population in Chihuahua, Mexico. A total of 6154 mistletoe plants were sexed on a total of 38 host trees; 3096 of these were male plants (50.3%) and 3058 were female (49.7%). A chi-square analysis indicated that these numbers were not significantly different from an essentially 1:1 sex ratio. Furthermore, although there was variation between trees (more male or female plants) the sex ratio on individual trees, and for each population, was also essentially 1:1. Key Words: Chihuahua pine dwarf mistletoe, Chihuahua pine, sex ratio Dwarf mistletoes (Arceuthobium spp., Visca- ceae) are dioecious, parasitic flowering plants that commonly occur on members of the Pinaceae in western North America. In general, the sex ratio (female plants : male plants) of dwarf mistletoes has been reported to be 1:1 (Hawksworth and Wiens 1996; Mathiasen et al. 1998). However, other in- vestigators have reported female-biased sex ratios for several dwarf mistletoes: Arceuthobium ameri- canum Engelm. in Canada (Muir 1966), A. globos- um Hawksw. & Wiens subsp. globosum and A. strictum Hawksw. & Wiens in Durango, Mexico (Hawksworth and Wiens 1996), A. pusillum Peck in Minnestoa (Baker et al. 1981), and most notably, A. tsugense (Rosendahl) G. N. Jones subsp. tsu- gense and subsp. mertensianae Hawksw. & Nick- rent from the Pacific Northwest and Alaska (Wiens et al. 1996). The majority of these reports of fe- male-biased sex ratios for dwarf mistletoes were based on small sample sizes (approximately 50— 500 total plants). However, the latter study used over 3000 plants from 16 populations distributed from southern Oregon to southeast Alaska and re- ported a significant female-biased sex ratio for 10 of the populations and for the pooled data from all 16 areas (59% females: 41% males). Because of the female-biased sex ratio reported for Arceuthobium tsugense (Wiens et al. 1996), Hawksworth and Wiens (1996) recommended that the sex ratios of other dwarf mistletoes be exam- ined. Therefore, this study was initiated to provide additional information on the adult sex ratios of dwarf mistletoes. Arceuthobium gillii Hawksw. & Wiens was chosen for this study because of the strong sexual dimorphism exhibited by this species (Hawksworth and Wiens 1996, see page 147). This strong sexual dimorphism allowed male and female plants to be easily distinguished. Other dwarf mis- tletoes also exhibit extreme sexual dimorphism and a study of this characteristic would be a valuable contribution to the systematics of the genus (Hawk- sworth and Wiens 1996, Mathiasen et al. 1998). MATERIALS AND METHODS The adult sex ratio for Arceuthobium gillii was determined for five populations in southern Arizona and for one population in Chihuahua, Mexico (Ta- ble 1, Fig. 1). At each of the locations sampled, at least four, severely infected Chihuahua pines (Pinus chihuahana Englem.) were selected, their diameters at breast height (1.3 m above the ground) measured to the nearest 0.5 cm, and a dwarf mistletoe rating (DMR) assigned to each tree using the 6-class sys- tem (Hawksworth 1977). For reasons of safety and efficiency sampling was restricted to trees <30 cm in diameter at breast height and a DMR >4. Selected trees were flagged and after diameters and dwarf mistletoe ratings were recorded, each tree was cut as close to the ground as possible. After each tree was cut, its total height to the near- est 0.1 m, was measured. Live branches that broke from the tree when it was cut were examined for mistletoe plants first. Then each live branch still attached to the tree, starting at the bottom of the crown, was removed and examined for mistletoe plants. The sex of each observed plant on a branch was recorded and the branch discarded well away from the tree so the branch would not be re-sam- pled. This process was repeated until all the live branches on each tree had been examined. Only plants that could be accurately sexed were tallied. One person examined branches for mistletoe plants and one person recorded data. This was to insure that the person examining mistletoe plants did not know how many males or females had been tallied as sampling proceeded. The same person recorded data until a tree had been completed. Data were not summarized until all the selected trees had been 2002] MATHIASEN AND DAUGHERTY: DWARF MISTLETOW SEX RATIO 13 TABLE 1. POPULATION LOCATIONS, NUMBER OF TREES SAMPLED, MEAN DIAMETERS, MEAN DWARF MISTLETOE RATINGS, MEAN TREE HEIGHTS, NUMBER OF PLANTS SAMPLED, AND ADULT SEX RATIOS (PERCENT FEMALE) FOR ARCEUTHOBIUM GILLI. No sex ratios exhibited a significant sex bias. Chi-square statistics (P = 0.05). DBH = diameter breast height; DMR = dwarf mistletoe rating (Hawksworth 1977). Mean Mean Trees DBH Mean height Plants Percent Population location sampled (cm) DMR (m) sampled female P Bear Canyon Santa Catalina Mountains, AZ 7 16.0 Deo} 8.5 1094 47.9 0.164 Gardner Canyon Santa Rita Mountains, AZ 4 15.0 5.8) 7.0 Syl 50.5 0.825 Carr Canyon Huachuca Mountains, AZ 9 16.0 5.6 8.1 1348 50.6 0.663 Pinery Canyon Chiricahua Mountains, AZ 4 18.0 5.6 8.3 1936 50.7 0.525 Upper Cave Creek Chiricahua Mountains, AZ 7 20.0 5.6 10.6 568 49.3 OW3m Chihuahua, Mexico (Sierra Madre Occidental) 7 16.0 Sis) Woe 697 48.2 0.344 Total/mean 38 17.0 5.4 8.4 6154 49.7 0.702 120° ey 110° 1053 JOD Arizona 35° New Mexico 30° ie TOE 105° Fic. 1. Approximate locations of study sites. 1—Bear Canyon, Santa Catalina Mountains, Arizona; 2—Gardner Can- yon, Santa Rita Mountains, Arizona; 3—-Carr Canyon, Huachuca Mountains, Arizona; 4—Pinery Canyon, Chiricahua Mountains, Arizona; 5—Upper Cave Creek, Chiricahua Mountains, Arizona; and 6—Sierra Madre Occidental, Chi- huahua, Mexico. 14 MADRONO [Vol. 49 TABLE 2. ADULT SEX RATIOS (PERCENT FEMALE) FOR ARCEUTHOBIUM GILLII FOR EACH TREE SAMPLED IN CARR CANYON, HuAcHUCA MOUNTAINS, ARIZONA. No sex ratios exhibited a significant sex bias. Chi-square statistics (P = 0.05). DBH = diameter breast height; DMR = dwarf mistletoe rating (Hawksworth 1977). dire DBH Height Plants Percent number (cm) DMR (m) sampled female IP 1 16.0 5 8.1 93 54.8 0.351 yy 10.5 6 5.6 153 53.6 0.374 3 19.5 6 10.3 196 49.0 0.775 4 16.0 5 7.8 76 44.7 0.359 5) 18.0 5 8.7 141 S32 0.449 6 14.0 6 Wood) 83 48.2 0.742 7 20.5 6 8.6 268 49.3 0.807 8 16.5 6 8.1 164 47.6 L532 9 16.0 5 8.0 174 54.0 0.289 Total/mean 16.0 5.6 8.1 1348 50.6 0.663 sampled. After summarizing the total number of plants sexed for the selected trees, additional trees were sampled to bring the total number of plants sexed to a minimum of 500, if necessary. Data were collected in May of 1998—2000 and in June 2001. This was either during anthesis of male plants or shortly after it (June). A chi-square analysis was used to determine if the ratio of male to female mistletoe plants exhibited a sex bias. We used a P value of =0.05 to determine the existence of statistically significant differences (Zar 1999). RESULTS The general location of study sites, the number of trees sampled, tree mean diameters, mean dwarf mistletoe ratings, mean heights, the number of mis- tletoe plants sexed, and the percentage of female plants for each population sampled are presented in Table 1. We sampled a total of 6154 mistletoe plants that could be accurately sexed on 38 trees. Of the plants we sexed, 3096 (50.3%) were males and 3058 (49.7%) were females. The difference in the number of male and female plants was not sig- nificantly different from the number expected for a 1:1 sex ratio (P = 0.628); therefore, the adult sex ratio of Arceuthobium gillii on the 38 trees was essentially 1:1 (Table 1). There was a large amount of variation in the sex ratio for A. gillii between individual trees at each of the study sites. An example of this variation is illustrated by the trees sampled in the Huachuca (Table 2) and Santa Rita Mountains, Arizona (Table 3). Some trees had more female plants than males and vice versa, but no trees exhibited significantly different sex ratios from a 1:1 ratio. In addition, when the sex ratio was determined using all of the trees in each population, no significant differences from a 1:1 sex ratio were detected for any of the populations (Tables 1-3). DISCUSSION Although several investigators have reported sig- nificant female-biased adult sex ratios for several dwarf mistletoes, we found that the adult sex ratio for Arceuthobium gillii is essentially 1:1. These re- sults were expected because many other dwarf mis- tletoes also have 1:1 adult sex ratios (Hawksworth and Wiens 1996; Mathiasen et al. 1998). Because many of the reports of female-biased adult sex ra- tios for dwarf mistletoes are based on relatively small sample sizes (<500 plants), they may not rep- resent an accurate estimate of the sex ratios for these mistletoes (Mathiasen and Shaw 1998; Daugherty and Mathiasen 1999). Our results demonstrate the variation in sex ratio that can occur among individual trees. This tree-to- tree variation has been demonstrated in other stud- ies of mistletoe sex ratio (Nixon and Todzia 1985; Mathiasen and Shaw 1998; Daugherty and Mathi- asen 1999). Because of this tree-to-tree variation, a large sample of mistletoe plants should be sampled for dioecious mistletoe sex ratio studies and data TABLE 3. ADULT SEX RATIOS (PERCENT FEMALE) FOR ARCEUTHOBIUM GILLII FOR EACH TREE SAMPLED IN GARDNER CAN- YON, SANTA RITA MOUNTAINS, ARIZONA. No sex ratios exhibited a significant sex bias. Chi-square statistics (P = 0.05). DBH = diameter breast height; DMR = dwarf mistletoe rating (Hawksworth 1977). Tree DBH Height Plants Percent number (cm) DMR (m) sampled female iE 1 14.5 5 al ae) 50.5 0.925 2 10.0 6 Se 123 54.5 0.321 3 20.0 5 V8 206 49.0 0.781 4 16.0 5 7.8 69 49.3 0.904 Total/mean 15.0 53) 7.0 Sill 50.5 0.825 2002] MATHIASEN AND DAUGHERTY should be analyzed using the results for the entire population and not on an individual tree basis (Ma- thiasen and Shaw 1998). We sampled over 6000 adult plants from several separate populations of A. gillii and found that this species exhibits a 1:1 adult sex ratio in each population sampled and when the data were pooled for all populations. Therefore, we contend that A. gillii will exhibit a consistent 1:1 adult sex ratio whenever large numbers of plants (>500) are sexed for populations of this dwarf mis- tletoe. We plan to determine the adult sex ratios for oth- er dwarf mistletoes in the Southwest and for those already reported to have female-biased sex ratios based on small samples, such as Arceuthobium americanun, A. globosum, and A. strictum (Hawk- sworth and Wiens 1996). However, we now hy- pothesize that when a large number of dwarf mis- tletoe plants are sampled for each of these dwarf mistletoes, the overall adult sex ratio will be essen- tially 1:1 as it was for the populations of A. gillii we sampled in this study. ACKNOWLEDGMENTS We would like to extend our appreciation to Del Wiens for suggesting this study be conducted. The field assis- tance of Dave Russell in the Chiricahua Mountains is sin- cerely appreciated also. LITERATURE CITED BAKER, FE A., D. W. FRENCH, AND G. W. HUDLER. 1981. Development of Arceuthobium pusillum on black spruce. Forest Science 27:203-—205. MADRONO, Vol. 49, No. 1, p. 15, 2002 : DWARF MISTLETOW SEX RATIO 15 DAUGHERTY, C. M. AND R. L. MATHIASEN. 1999. Adult sex ratio of Phoradendron juniperinum in ten severely infected Juniperus monosperma in northern Arizona. Madrono 46:169—176. HAWKSWORTH, E G. 1977. The 6-class dwarf mistletoe rat- ing system. USDA Forest Service Research Note RM-48. AND D. WIENS. 1996. Dwarf mistletoes: biology, pathology, and systematics. USDA Forest Service Agric. Handb. 709. MATHIASEN, R. L., C. G. PARKS, B. W. GEILS, AND J. S. BEATTY. 1998. Notes on the distribution, host range, plant size, phenology, and sex ratio of two rare dwarf mistletoes from Central America: Arceuthobium hawksworthii and A. hondurense. Phytologia 84:154— 164. AND D. C. SHAw. 1998. Adult sex ratio of Arceu- thobium tsugense in six severely infected Tsuga het- erophylla. Madrono 45:210—214. Murr, J. A. 1968. Biology of dwarf mistletoe (Arceuthob- ium americanum) in Alberta. Internal Report A15, Canada Department of Fisheries and Forestry, pp. 1— 29, Calgary, Alberta. NIXoNn, K. C. AND C. A. TobziA. 1985. Within-population, within-host species, and within-host tree sex ratios in mistletoe (Phoradendron tomentosum) in central Tex- as. American Midland Naturalist 114:304—310. WIENS, D., D. L. NICKRENT, C. G. SHAW, E G. HAwK- SWORTH, P. E. HENNON, AND E. J. KING. 1996. Embry- onic and host-associated skewed adult sex ratios in dwarf mistletoe. Heredity 77:55—63. ZAR, J. H. 1999. Biostatistical analysis. Prentice Hall, Up- per Saddle River, NJ. ANNOUNCEMENT First ANNOUNCEMENT AND CALL FOR SESSIONS FOURTH INTERNATIONAL SYMPOSIUM ON GRASS SYSTEMATICS AND EVOLUTION AND THIRD INTERNATIONAL CONFERENCE ON THE COMPARATIVE BIOLOGY OF THE MONOCOTYLEDONS The Third International Conference on the Com- parative Biology of the Monocotyledons and Fourth International Symposium on Grass Systematics and Evolution will be hosted by Rancho Santa Ana Bo- tanic Garden (Claremont, CA, USA) on 30 March— 5 April 2003. Topics will include morphology, anatomy, development, reproductive biology, mo- lecular biology, cytology, genomics, genetics, bio- chemistry, paleobotany, phylogenetics, classifica- tion, biogeography, ecology, and data integration. Sessions will be devoted to particular groups within monocots such as grasses and orchids. Monocots III will provide a rare opportunity for researchers in diverse fields to interact, share ideas, and form collaborations. We invite proposals from those who wish to organize sessions. A call for contributed papers and posters will follow. The conference pro- ceedings will be published. Springtime marks the flowering peak of the diverse California flora, and field trips are planned. Visit www.monocots3.org for conference details; or write Monocots III, Ran- cho Santa Ana Botanic Garden, 1500 North College Avenue, Claremont, CA 91711-3157 USA; E-mail info@monocots3.org; fax 1.909.626.7670; tele- phone 1.909.625.8767 ext. 333. Co-sponsors in- clude the American Society of Plant Taxonomists, Botanical Society of America, and the International Association for Plant Taxonomy. MaproNno, Vol. 49, No. 1, pp. 16-19, 2002 ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE (POLYGONACEAB), A NEW VARIETY FROM THE SOUTHERN SIERRA NEVADA, CALIFORNIA DANA A. York! Death Valley National Park, PO. Box 579, Death Valley, CA 92328 ABSTRACT Eriogonum ovalifolium var. monarchense is a new variety discovered on a limestone formation in the southern Sierra Nevada. It is only known from one population in the Kings River drainage basin. It is morphologically similar to E. ovalifolium var. vineum; an endangered species found on limestone outcrops in the San Bernardino Mountains. The habit of the two varieties is different primarily in the angle that the flowering stems arise from the base. Key words: Eriogonum ovalifolium var. monarchense, Kings River, Sierra Nevada, Monarch buckwheat, limestone In 1995, botanical explorations in the Kings Riv- er canyon of the southern Sierra Nevada yielded three previously unknown vascular plant taxa from the limestone (marble) outcrops around Boyden Cavern, near Kings Canyon National Park. Heter- otheca monarchensis Semple, Shevock, & York and Gilia yorkii Shevock & A. G. Day were de- scribed within a few years of their discovery. A new variety of buckwheat from the Boyden Cavern limestone required years of research and follow-up collecting to verify its taxonomic status. The first collections of the three new taxa were made on the same day (31 July 1995) in Monarch Wilderness (Sierra and Sequoia National Forests). Eriogonum ovalifolium Nutt. var. monarchense D. A. York, var. nov. (Fig. 1)—Type: USA, CA, Fresno Co, 86 km E of Fresno, Sierra National Forest, Monarch Wilderness, 2.4 km NW of Boy- den Cave on N side of the Kings River canyon, 36°50'08"N, 118°49'19"W (NAD 83), 1815 m, 31 July 1995, York 111 & Shevock (holotype CAS; isotypes JEPS, NY). Paratype: USA, CA, Fresno Co, 86 km E of Fresno, Sierra National Forest, Monarch Wilderness, 2.4 km NW of Boyden Cave on N side of Kings River canyon, 36°50'08"N, 118°49'19"W (NAD 83), 1815 m, 19 July 1996, York 1250 (RSA). Eriogonum ovalifolium var. vineum accedentes sed caules floriferentes decumbenti ad ascendenti sunt. Pulvinate perennials forming mats up to 30 cm across (Fig. 1A); leaves basal, petiolate, tomentose, 5—22 mm long, the margins flat to slightly crisped, petioles 2-10 mm long, blades elliptic to orbicular, 3-12 mm long, 3-12 mm wide; flowering stems scapose, l|—many per matted clump, decumbent to ascending, 2—6(9) cm long, tomentose to floccose; 'R-mail: Dana_York @nps.gov inflorescences capitate, the head 1.5—4 cm across; bracts scale-like, 3, 1-5 mm long; involucres clus- tered 4—6 per head, sessile, tomentose, turbinate, 5—8 mm long, with 5 rounded or acute teeth up to 2 mm long; flowers white to cream with green (ag- ing red) midribs, 4-6 mm long, glabrous, the peri- anth lobes dimorphic, the perianth lobes of the out- er whorl mostly twice the width of the inner whorl (Fig. 1B); stamens mostly exserted, 1-3 mm long, the anthers 0.4—0.6 mm long; achenes brown, 2-3 mm long, glabrous. DISTRIBUTION, HABITAT, AND PHENOLOGY Eriogonum ovalifolium var. monarchense (Mon- arch buckwheat) is a rare neoendemic found in eastern Fresno County, in the southern Sierra Ne- vada. The only known population (type locality) grows on the north side of a limestone formation in the Kings River canyon above 1800 m (5900 feet), in the vicinity of Boyden Cavern. The plants are located in Monarch Wilderness, just below the Monarch Divide, in the Sierra National Forest. Monarch Divide rises over 880 m above the canyon floor and is the boundary between the middle and south forks of the Kings River and the Sierra and Sequoia National Forests. The divide is rugged and varied with a mix of metamorphic and igneous rocks. The limestone component is typified by steep slopes and sheer cliffs. Eriogonum ovalifolium var. monarchense forms dense mats on ledges and crevices in sandy soils developed from decomposed limestone. The popu- lation consists of approximately 30 plants scattered over a few thousand square meters. Flowers are present from June to August. Associates include Argyrochosma jonesii (Maxon) Windham, Bromus madritensis L. ssp. rubens (L.) Husn., Cercocarpus intricatus Wats., Erigeron aequifolius Hall, Erysi- mum capitatum (Dougl.) Greene ssp. capitatum, Garrya flavescens Wats., Gilia yorkii, Heuchera ru- bescens Torr. var. rydbergiana Rosend., Butt. & 2002] = ‘5 SS ae 6 D> Pk ae te ¢ - , 4.0. ede sexe Fic. 1. ex 2 x ; oye ISS. Se) br as S' YORK: ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE 1) 1 cm Habit of Eriogonum ovalifolium Nutt. var. monarchense D. A. York, and a detailed illustration of a flower from holotype collection and photographs. A. Mature plant in flower. B. Flower with detail of perianth lobes in an erect position. Drawings by Laura Cunningham. Lak., Pinus monophylla Torr. & Frém., Selaginella asprella Maxon, Streptanthus fenestratus (Greene) J.T. Howell, and Yucca whipplei Torr. RELATIONSHIPS The new taxon differs in several respects from the other California varieties of Eriogonum ovali- folium (Table 1). Eriogonum ovalifolium var. mon- archense is morphologically similar to E. ovalifol- tum Nutt. var. vineum (Small) Jepson; an endan- gered species known only from limestone outcrops in the San Bernardino Mountains. Eriogonum oval- iYfolium var. vineum has flowering stems that are generally erect in contrast to the decumbent to as- cending flowering stems of E. ovalifolium var. mon- archense. These two varieties occur in disjunct mountain ranges and are approximately 325 km apart from each other. Eriogonum ovalifolium Nutt. var. purpureum (A. Nelson) Durand is a variety that occurs in the east- ern Sierra Nevada and other mountain ranges in western North America on various substrates (Re- veal 1989). It differs from E. ovalifolium var. mon- archense by having longer leaf blades and shorter involucres. Eriogonum ovalifolium Nutt. var. nivale (Canby) M.E. Jones is geographically close to, but not sym- patric with, E. ovalifolium var. monarchense. In California, it occurs mostly on granites in subalpine and alpine habitats throughout the Sierra Nevada, Cascades, and White Mountains (Hickman 1993). It differs from E. ovalifolium var. monarchense by having generally smaller leaves and shorter flow- ering stems and perianth lobes. KEY TO THE CALIFORNIA VARIETIES OF ERIOGONUM OVALIFOLIUM las-Blowers yellow. sm << 24a. . 2.2 var. ovalifolium 1b. Flowers white, cream, red, or purplish 2a. Flowers 2—3 mm long var. nivale 18 COMPARISON OF ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE WITH THE OTHER E. OVALIFOLIUM VARIETIES KNOWN FROM CALIFORNIA. TABLE 1. E. o. vineum E. o. purpureum E. o. ovalifolium E. o. nivale E. o. eximium E. o. monarchense Characters erect erect erect erect erect decumbent to ascend- Flowering stems ing 2-5 cm long unmargined 3-6 cm long unmargined orbicular 5—20 cm long + unmargined obovate 4—20 cm long + unmargined obovate 0.3—5 cm long 5—10 cm long + unmargined orbicular brown-margined Leaf blades elliptic to spatulate 0.5—2 cm long 4—6.5 mm elliptic to orbicular 0.3—1.2 cm long 5-8 mm 0.6—1.2 cm long 5—7 mm 0.5—2 cm long 4—6.5 mm 1-6 cm long 4—6.5 mm yellow 0.2—0.8 cm long 2—4 mm Involucre lengths Flowers white to cream purple or white to white to cream white to cream white to cream cream 4—6 mm long 3-5 mm long rocky 2-3 mm long 4—6 mm long 4—6 mm long sandy granites 4—6 mm long rocky sandy or gravelly various substrates W. North America sandy or gravelly various substrates W. North America sandy or gravelly granites Habitat limestone limestone San Bernardino Mtns. W. North America N. Sierra Nevada, S. Sierra Nevada, Range Nevada 1800—3400 m elev. California 1800 m elev. MADRONO 1200—2800 m elev. 1500—2100 m elev. 1200—2900 m elev. 1500—4100 m elev. [Vol. 49 2b. Flowers >3 mm long 3a. Leaves obviously brown-margined Ce nr Cae. Meet, AS a Sic var. eximium 3b. Leaves not distinctly brown-margined 4a. Flowering stems 4—20 mm long; in- volucres 4—6.5 mm long; various SUDSERateSs eee es ee var. purpureum 4b. Flowering stems 2—6 (9) mm long; involucres 5—8 mm long; carbonate substrates 5a. Flowering stems decumbent to ascending; southern Sierra Ne- vada Range var. monarchense 5b. Flowering stems mostly erect; San Bernardino Mountains .. . A AE rte S25 5 var. vineum DISCUSSION Eriogonum ovalifolium var. monarchense is al- lopatric from the other varieties of E. ovalifolium. The calcareous habitat and associated species where Eriogonum ovalifolium var. monarchense is found are more typical of pinyon pine communities in the eastern Sierra Nevada and the desert ranges of California and Nevada. It is possible that Erio- gonum ovalifolium var. monarchense is allied with the varieties from the desert and southern Califor- nia mountains and not with the Sierran Eriogonum ovalifolium var. nivale. There are four other calci- cole vascular plants endemic to the King River ba- sin (York 1999). These include Eriogonum nudum Benth. var. regirivum Reveal & J. Stebbins, Gilia yorkil, Heterotheca monarchensis, and Streptanthus fenestratus. Heterotheca monarchensis (occurs on south-facing slopes with a population very near Er- togonum ovalifolium var. monarchense) and Gilia yorkii are allied with their respective desert con- geners (York 1999). They evolved from desert taxa that spread into the California Floristic “Province during Xerothermic periods of the Quaternary (York 1999). Other evidence of this desert link in- clude taxa found on the limestone outcrops around Boyden Cavern, such as Achnatherum hymenoides (Roem. & Schultes) Barkworth, Argyrochosma jo- nesii, Melica frutescens Scribn., and Petrophyton caespitosum (Nutt.) Rydb., that are mostly rare in the Sierra Nevada and common in the desert ranges (York 1999). RARITY Eriogonum ovalifolium var. monarchense, pre- viously unknown and uncollected, is a rare taxon due to its lithophytic nature on limestone, a rel- atively uncommon substrate in the southern Si- erra Nevada. Because of the remoteness and rug- ged physiography of the Kings River canyon limestones, it is unlikely that Eriogonum ovali- folium var. monarchense occurs outside the river basin. It is a rare and localized endemic worthy of conservation efforts. Anthropogenic impacts are not likely because the population is remote and within a designated wilderness. If the type 2002] locality is indeed the only population, then this taxon is vulnerable to extinction due to stochastic events or genetic drift. ACKNOWLEDGMENTS I thank Laura Cunningham for her carefully detailed work on the illustrations, and Johanna Arnegger (my Aus- trian friend) for correcting the Latin diagnosis. I also thank Jim Shevock for following me through poison oak, dense brush, loose rock, and up steep slopes during an uncom- fortably hot day to assist me in collecting the type mate- YORK: ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE 19 rial. James Reveal and an anonymous reviewer provided useful comments on my drafts for which I am grateful. LITERATURE CITED HICKMAN, J. C. 1993. Polygonaceae in J. Hickman (ed.), The Jepson manual: Higher plants of California. Uni- versity of California Press, Berkeley. REVEAL, J. L. 1989. The eriogonoid flora of California (Polygonaceae: Eriogonoideae). Phytologia 66:295— 414. York, D. A. 1999. A phytogeographic analysis of the Kings River Basin, California. M.S. thesis. California State University, Fresno, CA. MADRONO, Vol. 49, No. 1, pp. 20—21, 2002 HESPEROYUCCA WHIPPLEI AND YUCCA WHIPPLEI (AGAVACEAE) JEFFREY A. GREENHOUSE Jepson Herbarium, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, CA 94720-2465 JOHN L. STROTHER University Herbarium, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, CA 94720-2465 ABSTRACT The name Yucca whipplei dates from 1861, not 1859. The names Hesperoyucca, Hesperoyucca whip- plei, and Hesperoyucca whipplei var. graminifolia date from 1893. Chaparral yucca has been widely known for 140 years or so as Yucca whipplei. The plants have long been recognized as markedly unlike other yuccas (e.g., Engelmann 1871). Baker (1892) suggested the plants are generically distinct from other yuc- cas. But, chaparral yucca has continued to reside within Yucca in most taxonomic and floristic treat- ments. Primarily on molecular similarities and dif- ferences, Bogler and Simpson (1995), Clary and Simpson (1995), and others cited by them, have revived the suggestion that chaparral yucca should be placed in a separate genus, Hesperoyucca. Here, we review the histories of the botanical names. When the name Yucca whipplei was first used in print, Torrey (1859, p. 222) described (or at least diagnosed) the taxon and wrote, “If it prove to be a distinct species it may be called Y. Whipplei.”’ We consider that use of Yucca whipplei by Torrey to have been as a provisional name and, therefore, we believe that the name was not validly published in 1859. The next use of Yucca whipplei that we have found was also by Torrey (1861). Conditions for valid publication of Yucca whipplei were met by Torrey in 1861. We believe the correct citation to be: Yucca whipplei Torrey in J. C. Ives, Rep. Colorado R. 4(Botany):29. 1861. Plants of Yucca whipplei have also been called Hesperoyucca whipplei. There is no entry for Hes- peroyucca in the published version of ING (Farr et al. 1979). In June 2001, requests to the website for Index Nominum Genericorum (currently, http:// rathbun.si.edu/botany/ing/) for Hesperoyucca elic- ited the response, ““No records found with: Search for Name “‘Hesperoyucca.’ ”’ The earliest use of Hesperoyucca of which we are aware was by Engelmann (1871), who parti- tioned Yucca into: “§ EU-YUCCA,”’ with three subordinate groups (Sarcocarpa, Clistocarpa, and Chaenocarpa) with one species each, and *‘§ HES- PERO-YUCCA,” which included Yucca whipplei as the only species. In 1873, Engelmann provided a summary “SYSTEMATIC ARRANGEMENT” for Yucca in which he named: I. Euyucca (which included: A. Sarcoyucca, B. Clistoyucca, and C. Chaenoyucca) and II. Hesperoyucca. In both clas- sifications, the rankless Hesperoyucca included only Yucca whipplei. In a third paper (“‘corrections and additions” to the 1873 paper), Engelmann (1875) provided yet another “‘synopsis”’ for Yucca and that time parti- tioned Yucca into four codrdinate elements named Sarcoyucca, Clistoyucca, Chaenoyucca, and Hes- peroyucca. Once again, Engelmann did not indicate rank for his subdivisions of Yucca. Baker (1876), in notes on Yucca whipplei, wrote, **... Dr. Engelmann, who has considered it [i.e., Yucca whipplei|] as the type of a new subgenus, which, alluding to its western locality, he has called Hesperoyucca. At present this subgenus is only known to contain a single species.’’ We believe that Baker’s remarks were sufficient to establish sub- generic rank and that the correct citation is: Yucca L. subg. Hesperoyucca (Engelmann) Baker, Gard. Chron. n.s. 6:196. 1876. Based on Yucca [unranked] Hesperoyucca Engelmann in S. Watson et al., Botany (Fortieth Parallel), 497. 1871, as ““§ 2. HESPERO-YUCCA.” In 1892, Baker wrote of Y. whipplei, “‘I now think this had better be kept as a genus distinct from Yucca, under Engelmann’s name Hesperoyuc- ca.”’ Baker, nevertheless, treated whipplei as be- longing to Yucca in 1892. We suggest that Baker’s statement in 1892 may have been a “‘note-added- in-proof”’ and that Baker’s statement was not suf- ficient to establish generic rank for Hesperoyucca. Trelease (1893) accepted Baker’s suggestion of generic rank for Hesperoyucca, made a specific combination in Hesperoyucca, and coined a varietal nomen novum in Hesperoyucca. We believe correct citations for the names are: 2002] Hesperoyucca (Engelmann) Trelease, Rep. (An- nual) Missouri Bot. Gard. 4:208. 1893. Based on: Yucca [unranked] Hesperoyucca Engel- mann in S. Watson et al., Botany (Fortieth Par- allel) 497. 1871. —Type: Hesperoyucca whip- plei (Torrey) Trelease = Yucca whipplei Tor- rey. Hesperoyucca whipplei (Torrey) Trelease, Rep. (Annual) Missouri Bot. Gard. 4:208. 1893, as ‘Y. Whipplei’ under ‘HESPEROYUCCA’. Given the use of Hesperoyucca whipplei else- where in the article by Trelease (figure cap- tions, in a name at varietal rank), we consider the ““Y”’ at the comb. nov. to be a typographic error and the attribution of the combination to Baker to have been superfluous. Basionym: Yucca whipplei Torrey. Hesperoyucca whipplei (Torrey) Trelease var. gra- minifolia Trelease, Rep. (Annual) Missouri Bot. Gard. 4:215. 1893. We maintain that Tre- lease coined a nom. nov. based on: Yucca gra- minifolia Wood, 1868 [Proc. Acad. Nat. Sci. Philadelphia 20:167], non Zucc., 1837. After submitting this paper to Madrono for pub- lication (June 2001), drafts of the manuscript (or relevant excerpts) were sent to colleagues with es- pecial interest in botanical nomenclature. A flurry of e-mails ensued. Some nomenclaturists held that Baker validated Hesperoyucca at generic rank in 1892. Others opined that Baker did not validly pub- lish Hesperoyucca at generic rank and that Hespe- royucca was validly published at generic rank by Trelease in 1893. To us, Baker explicitly treated Hesperoyucca at subgeneric rank in 1876 and did not explicitly treat Hesperoyucca at generic rank in 1892. As a result of having the matter of valid publi- cation of Hesperoyucca at generic rank bandied about, the website for ING posted this entry in No- vember 2001: Hesperoyucca (G. Engelmann) J. G. Baker, Bull. Misc. Inform. Kew 1892: 8. Jan 1892. T.: H. whipplei (J. Torrey) W. Trelease (‘Y. whip- ple?) (Rep. (Annual) Missouri Bot. Gard. 4: 208. 1893) (Yucca whipplei J. Torrey) Yucca [unranked] Hesperoyucca G. Engelmann in S. Watson, U.S. Geol. Surv. 40th Parallel, Bot. 497. Sep—Dec 1871. PHAN.-AGAVACEAE (104) 06 Nov 2001 GREENHOUSE AND STROTHER: HESPEROYUCCA 21 In December 2001, after further bandying, the ING website entry was changed to: Hesperoyucca (G. Engelmann) W. Trelease, Rep. (Annual) Missouri Bot. Gard. 4: 208. 9 Mar 1893. T.: H. whipplei (J. Torrey) W. Trelease (‘Y. whip- pler) (Yucca whipplei J. Torrey) Yucca [unranked] Hesperoyucca G. Engelmann in S. Watson, U.S. Geol. Surv. 40th Parallel, Bot. 497. Sep-Dec 1871._For the publication date, see I.c. (Rep.) 5: 3. 1894. PHAN.-AGAVACEAE (104) 14 Dec 2001 ACKNOWLEDGMENTS This study resulted from work on the Jepson Flora Pro- ject and was supported in part by a grant from the William R. Hewlett Revocable Trust. We appreciate comments and opinions sent to us by e-mail from farflung colleagues. We thank D. J. Bogler and S. Verhoek for formal reviews of the manuscript and K. H. Clary for informal comments on an early version of this paper. LITERATURE CITED BAKER, J. G. 1876. New garden plants. Gard. Chron. n.s. 6:196—197. . 1892. CCXXIII-Agaves and arborescent Lili- aceae on the Riviera. Bull. Misc. Inform. [Kew] 61: 5-10. BOoGLER, D. J. AND B. B. Simpson. 1995. A chloroplast DNA study of the Agavaceae. Syst. Bot. 20:191—205. CLARY, K. H. AND B. B. Stmpson. 1995. Systematics and character evolution of the genus Yucca L. (Agava- ceae): Evidence from morphology and molecular analyses. Bol. Soc. Bot. México 56:77-88. ENGELMANN, G. 1871. Appendix (pp. 496—497) to S. Wat- son, List [Catalogue] of plants collected in Nevada and Utah, in S. Watson et al., Botany (Fortieth Par- allel). . 1873. Notes on the genus Yucca. Trans. St. Louis Acad. Sci. 3:17—54. . 1875. Notes on the genus Yucca. No. 2. Trans. St. Louis Acad. Sci. 3:210—214. FarR, E. R., J. A. LEUSSINK, AND E A. STAFLEU. 1979. Index nominum genericorum (plantarum), Vol. 2, Eprolithus—Peersia. Regnum Veg. 101:631—1276. TORREY, J. 1859 [1858]. Jn W. H. Emory, Report on the United States and Mexican boundary survey, Vol. 2, pp. [27]—270, Botany of the boundary. . 1861. In J. C. Ives, Report upon the Colorado River, part 4, pp. [1]—30, Catalogue of the plants col- lected upon the expedition. TRELEASE, W. 1893. Further studies of yuccas and their pollination. Rep. (Annual) Missouri Bot. Gard. 4: 181-226. MApRONO, Vol. 49, No. 1, pp. 22-24, 2002 SYMPATRY BETWEEN DESERT MALLOW, EREMALCHE EXILIS AND KERN MALLOW, E. KERNENSIS (MALVACEAE): MOLECULAR AND MORPHOLOGICAL PERSPECTIVES KATARINA ANDREASEN Laboratory of Molecular Systematics, Swedish Museum of Natural History, Stockholm, Sweden; Jepson Herbarium and Department of Integrative Biology, University of California, Berkeley CA 94720-2465 ELLEN A. CYPHER Endangered Species Recovery Program, 1900 North Gateway Boulevard, Suite 101, Fresno, CA 93727 BRUCE G. BALDWIN Jepson Herbarium and Department of Integrative Biology, University of California, Berkeley CA 94720-2465 ABSTRACT Molecular and morphological data support an extended distribution of Eremalche exilis into the south- ern San Joaquin Valley and southern Inner South Coast Ranges of California, within the range of the morphologically similar, rare, and endangered E. kernensis. Nuclear rDNA sequences for plants from Kern County that were morphologically indistinguishable from E. exilis were identical at all non-poly- morphic sites to sequences of undisputed EF. exilis from the Mojave Desert and Sonoran Desert, but differed from sequences of E. kernensis by seven sequence mutations. Morphologically, E. exilis can be distinguished from E. kernensis by anther position—in flowers of EF. exilis, the anthers are held at the same level as the stigmas; in bisexual flowers of E. kernensis, the anthers are situated well below the stigmas. Based on limited rDNA data, we found no evidence of gene flow between sympatric populations of E. exilis and E. kernensis. Key words: Eremalche, Malvaceae, California floristics, ITS, ETS, rare and endangered plants. Botanists have disagreed about whether the de- sert mallow, Eremalche exilis (A. Gray) Greene, occurs within the California Floristic Province (Kearney 1956; Twisselmann 1956, 1967; Hoover 1970; Bates 1992). In The Jepson Manual, Bates (1993) indicated that the distribution of E. exilis in California is restricted to the Mojave and Sonoran Deserts, whereas Twisselmann (1956, 1967) and Hoover (1970) indicated that £. exilis occurs in the San Joaquin Valley and Inner South Coast Ranges of western Kern County and eastern San Luis Obis- po County, California. If E. exilis is present in the southern San Joaquin Valley and southern Inner South Coast Ranges of California, then it may be of importance for evaluating the origin and status of the rare and endangered Kern mallow, E. ker- nensis C. B. Wolfe [=E. parryi subsp. kernensis (C. B. Wolfe) D. M. Bates pro parte; Bates (1992) ex- panded the circumscription of Kern mallow], which is endemic to the region. Eremalche kernensis has been suggested to be of hybrid origin between E. exilis and the widespread central Californian taxon E. parryi (Greene) Greene (Kearney 1956). A co- occurrence of these taxa would be relevant to that hypothesis. Here we present molecular and mor- phological data that support an extended distribu- tion of E. exilis into the southern San Joaquin Val- ley and southern Inner South Coast Ranges of Cal- ifornia, within the range of E. kernensis. MOLECULAR DATA Plants from Kern County that we could not dis- tinguish morphologically from E. exilis were in- cluded in an ongoing project that aims to clarify whether E. kernensis is evolutionarily distinct from E. parryi (i.e., that the two taxa are independent, non-interbreeding, evolutionary lineages). Individ- uals from two populations of putative E. exilis from western Kern County were sampled for the molec- ular analysis. One specimen (Cypher 99-1A) was collected in the Lokern Natural Area (T29S, R22E, sect. 29 SE% of NW% of SE%, MDM), and was growing intermixed with the rare and endangered Kern mallow, E. kernensis. The other specimen (Vanherweg 99-14) was collected on a hillside west of Derby Acres (T31S, R22E, sect. 10 E% of NW%, MDM). In addition, one sample of E. exilis from the Mojave Desert (Heckard 4508) as well as sey- eral samples of E. kernensis were included. The DNA was extracted from pressed and dried leaf material of individual plants and the Internal and 3'-External Transcribed Spacers (ITS and ETS) in 2002] ANDREASEN ET AL.: SYMPATRY OF EREMALCHE EXILIS AND E. KERNENSIS 23 TABLE 1 WARIABLE NUCLEOTIDE POSITIONS IN THE INTERNAL (ITS) AND EXTERNAL (ETS) TRANSCRIBED SPACER REGIONS OF E. EXILIS AND E. KERNENSIS. Vouchers are deposited at JEPS and sequences at EMBL. EC = E. Cypher; BV = B. Vanherweg; SB = San Bernardino Co.; * = non-identical nucleotide positions between E. exilis and E. kernensis; Each number corresponds to one DNA position and polymorphic positions are abbreviated: s = cg; y = ct; w = at; k = gt; r = ag; m = ac. ? = not sequenced. — = gap; NN = ac/-. Taxon Collection data Accession numbers ITS (1-646) and ETS (710-1152) positions ITS/ETS hes a3 a tah cat eee lL al alicia Galea aL 17111111711222444667778889999900001111 QL Da sy7/ 23S SSO IL 3 213} 7) 7/ QOL AG 10S} 335) 91205885908487260955567839462153782 E. exilis Kern EC 99-1A AJ416060/AJ4 16065 eget CGECagebacgeqegqgcEgtecEEGcrtacs E. exilis Kern BV 99-]4 AJ416062/AJ4 16067 cgtcctcagctacgceg?gggctgtccttycwtacs E. exilis SB Heckard 4508 AJ416061/AJ416066 catcctyagctacgmgmgkrctrtcmttyctyacg E. kernensis Kern EC 99-3 AJ416063/AJ416064 yoyyyycmrmeryrcc ?ngg--gctcwgcgttNNg the nuclear ribosomal DNA were amplified and se- quenced as described by Andreasen and Baldwin (2001). ITS and ETS sequences for plants identified on the basis of morphology as E. exilis from western Kern County were identical to the ITS and ETS sequences for indisputed E. exilis from the Mojave Desert, except at twelve nucleotide positions (Table 1). At eleven of these positions one or two of the E. exilis samples were polymorphic, with one ad- ditional nucleotide besides the one present in the non-polymorphic sample(s). At one position, E. ex- ilis from the Mojave Desert had a unique state (po- sition 111). ETS and ITS sequences of Eremalche kernensis (one sample sequence is shown in Table 1) were polymorphic for many positions but not for the same positions that were polymorphic in the samples of E. exilis. In addition to the polymorphic positions, FE. exilis samples and E. kernensis dif- fered at eight nucleotide positions in the ETS and ITS sequences. In the ITS region, the two taxa dif- fered by two point mutations (positions 268 and 646). In the ETS, E. exilis and E. kernensis differed by four point mutations (positions 938, 973, 1006, and 1021) and E. exilis was marked by a two base pair insertion (at positions 825 and 826). This in- Sertion was unique for E. exilis and was not present in E. rotundifolia or any other sequenced taxon from tribe Malveae (Andreasen and Baldwin 2001, unpublished data). In addition, ETS sequences of E. kernensis were polymorphic for a two base-pair indel (ac/-; NN in Table 1) at positions 1137 and 1138. All other Malveae sequences had ‘“‘ac”’ at these positions. Phylogenetic analysis of ETS and ITS sequences for each of the taxa of Eremalche and various outgroup taxa in tribe Malveae af- firmed that samples of E. exilis from the Mojave Desert and western Kern County constitute a line- age (data not shown). MORPHOLOGICAL AND DISTRIBUTIONAL DATA Overlapping variation in flower color, flower size, and growth form has contributed to uncertain- ty about the identity of Eremalche populations in the southern San Joaquin Valley and southern Inner South Coast Ranges. Both E. exilis and E. kernensis most commonly have white petals, although petals of E. exilis may be tinged with pink and petals of E. kernensis occasionally are pale lavender (Wolf 1938; Kearney 1956; Cypher unpublished data). Keys to species of Eremalche typically have used petal and calyx lengths to distinguish E. exilis from E. kernensis and E. parryi (e.g., Kearney 1956; Munz 1968; Bates 1993). However, these keys failed to differentiate petal length by gender. Ere- malche exilis has only bisexual flowers, whereas E. kernensis is gynodioecious, 1.e., with bisexual-flow- ered plants and pistillate-flowered plants (Bates 1992, 1993). In E. kernensis, petals of pistillate plants are shorter (3 to 7 mm long) than petals of bisexual plants (4.5 to 10 mm long) (Cypher un- published data). Pistillate plants of E. kernensis have petals that often are similar in size to those of E. exilis, which has petals ranging in length from 4 to 5.5 mm (Bates 1993). For the most part, pistillate flowers of E. kernen- sis have only styles and no stamens, but in a small percentage of flowers, there may be a few (less than five, Cypher unpublished data) vestigial stamens that are reduced in size and do not shed pollen. The absence of stamens (or, more infrequently, the pres- ence of few, vestigial, non-functional stamens) in small-flowered (pistillate) plants of E. kernensis should be sufficient to distinguish morphologically E. kernensis from E. exilis, but the differing posi- tion of the anthers relative to the stigmas in the two taxa has caused confusion. In £. exilis, the anthers are held at the same level as the stigmas, whereas in bisexual flowers of E. kernensis the anthers are situated well below the stigmas, near the base of the corolla. Thus, bisexual flowers of E. exilis have been mistaken for pistillate flowers of E. kernensis, when anthers at the same level as the stigmas have been interpreted as stigmas. The growth form of E. exilis has been described as prostrate or decumbent (Munz 1968; Bates 1993), whereas Eremalche ker- 24 MADRONO nensis stems may be erect or prostrate (Wolf 1938; Munz 1968). We have observed erect, slender- stemmed plants of E. exilis in western Kern Coun- ty, as well as individuals of both E. exilis and E. kernensis that have a stout, erect central stem and several prostrate lateral stems. Investigations in the Lokern area of western Kern County (west of the community of Button- willow) from 1996 through 2001 revealed that E. kernensis does not range more than 3.2 km south of Lokern Road. The southernmost occurrences of E. kernensis that we have confirmed are in T29S, R22E, sect. 27, 28, and 29, MDM. Based on motr- phological characters, we have confirmed the pres- ence of E. exilis in T29S, R22E, sect. 14, 19, 20, a), Pe, Dy 2S, 2, SZ, BG! BS, IMIDE im WIOS, R22E, sect. 4; and in T31S, R22E, sect. 10, MDM. Eremalche exilis and E. kernensis are sympatric in at least T29S, R22E, sect. 14, 20, 23, 27, 28, and 29, MDM, sometimes growing in mixed colonies. In addition to the above mentioned localities we have identified four collections (at UC) from the southern Inner South Coast Ranges and southern San Joaquin Valley of California that match the de- scription of E. exilis: (1) Hoover 9350, collected in 1965 in San Luis Obispo County (“Hill just east of San Juan River, La Panza District, in sandy calcar- eous soil’’); (2) Hoover 441, collected in 1935 in Fresno County (“10 mi e. of Coalinga’’); (3) Alice Eastwood s. n., from “Huron, Calif.’’ in Fresno County (this collection is from 1893, before the Huron vicinity was converted to farmland); and (4) Dean Wm. Taylor 10171, collected in 1989 in Kern County (“‘just west of California Aqueduct ca. 1.3 miles SE of Highway 58’’). The fourth collection (Dean Wm. Taylor 10171) yielded ETS and ITS sequences that were identical to ETS and ITS se- quences of one of the FE. exilis specimens from Kern County described above (Cypher 99-1A) ex- cept at ETS position 1152, where DWT 1/0171] had a “g’ and Cypher 99-1A had a “cg.” CONCLUSIONS The morphological and molecular data presented here lead us to suggest that Eremalche exilis has a wider distribution in California than the distribution reported in The Jepson Manual (Bates 1993). The distribution of E. exilis should be extended beyond the Mojave Desert and Sonoran Desert to include the California Floristic Province (southern San Joa- quin Valley and Inner South Coast Ranges) in Kern and San Luis Obispo counties. Since there are no known recent collections of E. exilis from Fresno County it is uncertain if it still occurs there. The [Vol. 49 habitat destruction in the San Joaquin Valley sug- gests that E. exilis is likely to have disappeared from the Huron area. In contrast to the extended distribution of EF. exilis, the range of the rare and endangered FE. kernensis may be even narrower than previously believed because of earlier mis- identifications of E. exilis as E. kernensis. The molecular data demonstrate that E. exilis and E. kernensis are genetically and evolutionarily dis- tinct entities. Seven sequence differences between them were found, including one insertion unique for E. exilis. These positions are not polymorphic in either taxon, contrary to the expectation for hy- bridizing taxa. We found no evidence of gene flow between E. exilis and E. kernensis in rDNA se- quences of plants collected from the one area of sympatry sampled in this study. The previously proposed hybrid origin of E. kernensis and its ge- netic distinctness from EF. parryi is under continu- ing investigation and will be addressed in a later paper. ACKNOWLEDGMENTS We thank B. Vanherweg for collecting one of the Ere- malche exilis specimens included in this investigation. We also thank B. Wessa for DNA sequencing of one of the E. exilis samples (DWT 10171). This study was financed by a grant from the US Fish and Wildlife Service for a biosystematic study of Kern mallow. LITERATURE CITED ANDREASEN, K. and B. G. BALDWIN. 2001. Unequal evo- lutionary rates between annual and perennial lineages of checker mallows (Sidalcea, Malvaceae): Evidence from 18S—26S rDNA internal and external tran- scribed spacers. Molecular Biology and Evolution 18: 936-944. Bates, D. M. 1992. Gynodioecy, endangerment, and sta- tus of Eremalche kernensis (Malvaceae). Phytologia 72:48—54. . 1993. Eremalche. P. 748 in J. C. Hickman (ed.), The Jepson manual: Higher plants of California. Uni- versity of California Press, Berkeley. Hoover, R. FE 1970. The vascular plants of San Luis Obis- po County, California. University of California Press, Berkeley. KEARNEY, T. H. 1956. Notes on Malvaceae. VIII. Ere- malche. Madrofio 13:241—243. Munz, P. A. 1968. A California flora and supplement. Uni- versity of California Press, Berkeley. TWISSELMANN, E. C. 1956. A flora of the Temblor Range and the neighboring part of the San Joaquin Valley. The Wasmann Journal of Biology 14:161—300. . 1967. A flora of Kern County, California. The Wasmann Journal of Biology 25:1—393. WoLF, C. B. 1938. California plant notes II. Occasional Papers Rancho Santa Ana Botanic Garden, Series I, 2:44—-90. MaproNo, Vol. 49, No. 1, pp. 25-32, 2002 POLLINATION OF CYTISUS SCOPARIUS (FABACEAE) AND GENISTA MONSPESSULANA (FABACEAE), TWO INVASIVE SHRUBS IN CALIFORNIA INGRID M. PARKER! AND ALEXANDRA ENGEL Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, CA 95064 KAREN A. HAUBENSAK Department of Integrative Biology, University of California, Berkeley, CA 94720 KAREN GOODELL Department of Ecology, Evolution, and Natural Resources, Rutgers University, New Brunswick, NJ 08903 ABSTRACT Mutualistic interactions between natives and non-natives, and between different introduced species, can play an important role in the invasion process. The facilitation of a new introduced species by a previous invader could either accelerate an invasion or exacerbate its impact, providing a positive feedback loop in heavily invaded ecosystems. Open grasslands in Marin County, CA, are being invaded by two closely related, introduced legumes, Cytisus scoparius (Scotch broom) and Genista monspessulana (French broom). These non-clonal shrubs have been shown to be non-autogamous and pollen limited, underscoring the potential importance of pollinators to their fecundity and spread. The flowers of both are fused shut and require forced “tripping” by a pollinator. We measured floral characters and pollen production to make predictions about which species would be most attractive, and most accessible, to bee visitors. Cytisus flowers were an order of magnitude larger and produced four times as many pollen grains, suggesting that they should be more attractive and rewarding than Genista flowers. However, Cytisus flowers also required significantly more force to open, suggesting that less powerful pollinator species might be excluded from visiting. We tested these predictions by quantifying visitation rates and directly observing pollinators at two sites where the invaders co-occur. Consistent with the mechanical assay, pollinators were more successful at accessing flowers of the small-flowered Genista than the large-flow- ered Cytisus; however, Cytisus was more frequently visited than Genista, suggesting that pollinators preferred the larger and more rewarding species. We did not find evidence to support the prediction that the small-flowered Genista was accessible to a greater diversity of pollinator species. Although introduced from the same native range as the two plant invaders, honey bees (Apis mellifera) were not “‘better”’ pollinators than native bumble bees (Bombus vosnesenskii) in terms of effectiveness at tripping flowers or the number of flowers visited per plant. However, Apis was the numerically dominant pollinator at both sites, underscoring the potential conservation implications of local Apis introduction for the spread of noxious weeds in natural ecosystems. Key words: exotic, introduced, alien, mutualism, plant-pollinator, invasional meltdown INTRODUCTION Biological invasions have become one of our most alarming conservation issues (Williamson 1996; Parker and Reichard 1998; Mooney and Hobbs 2000). Negative interactions such as com- petition and predation between introduced and na- tive species (“‘biotic resistance’’) have long been thought to play a central role in determining wheth- er an invasion will succeed or fail (Darwin 1865; Elton 1958; Levine 2000; Maron and Vila 2001). However, mutualistic interactions between natives and non-natives may also be important in the in- vasion process (Simberloff and Von Holle 1999; Richardson et al. 2000). Interactions between dif- ‘Author to whom correspondence should be addressed. Email: parker @biology.ucsc.edu ferent introduced species can be important too, as suggested by the idea of “‘invasional meltdown” (Simberloff and Von Holle 1999; Parker et al. 1999), where the facilitation of a new introduced species by a previous invader could either accel- erate an invasion or exacerbate its impact, provid- ing a positive feedback loop in heavily invaded ecosystems. For example, spread of the nitrogen- fixing tree Myrica faya in Hawai’i appears to occur primarily through dispersal by an introduced bird species, Zosterops japonica (Vitousek and Walker 1989). The interaction between plants and their polli- nators is an example of a mutualism that can exert strong control over the fitness of the interacting partners and is thought to have driven the evolution of many floral traits (Darwin 1877; van der Pijl 1961). Pollinator limitation of plant reproduction is 26 MADRONO common (Burd 1994) and could be especially im- portant for introduced species. A successful inva- sive plant must be able to colonize new territory, usually in small numbers, and in the absence of its original pollinators. Such a scenario would appear to favor species that are self-fertile and capable of autonomous self-pollination, or autogamy (Allard 1965; Baker 1965; Brown and Marshall 1981; Bar- rett 2000). Open grasslands in Marin County, California, are being invaded by two closely related European shrubs, Cytisus scoparius (L.) Link (Scotch broom) and Genista monspessulana (L.) L. Johnson (French broom). Both species were first introduced to California as ornamentals in the 1860’s and 1870’s (Hoshovsky 1986). Cytisus and Genista are non-clonal legumes, are very similar in appearance, and are often lumped together in discussions of their ecological role as invaders (McClintock 1985; Hoshovsky 1986). Despite the above prediction that successful invaders should be self-fertile and au- togamous, in previous work we found that neither of these species showed substantial autogamy when pollinators were excluded, and that both exhibited reduced seed set when fertilized with self pollen (Parker and Haubensak 2002). We also found that both species were significantly pollen limited in populations on the Marin Peninsula (Parker and Haubensak 2002). These results demonstrate that the abundance and behavior of pollinators have im- portant consequences for the reproductive success of both these invaders. At the same time, however, there are apparent differences in floral biology be- tween the two species which we predicted could differentially affect their pollination success. Flow- ers of Cytisus appear larger than those of Genista. Reproductive plants of both species vary in size from 0.5 m to >2 m, so although Genista produces more of its small flowers for a branch of the same size (Parker and Haubensak, unpublished data), both plants can have from several dozen to several thousand yellow flowers. Therefore the primary dif- ference in floral display that distinguishes the spe- cies appears to be flower size. Neither plant pro- duces nectar, so pollen is the only reward for insect visitors. Theoretical expectations led us to pose two al- ternative hypotheses about how differences be- tween the two species could influence their repro- ductive success. The first hypothesis involves the ease of handling of flowers by pollinators. Larger flowers like those of Cytisus may be more difficult to handle, preventing the access of certain insect species and leading to specialization on a narrower set of pollinators. Both Cytisus and Genista have typical papilionaceous legume flowers, but with fused keel petals that an insect visitor must split in order to release the style and anthers. Flower visi- tors may have to apply more force to trip larger flowers (Westerkamp 1997), which could restrict [Vol. 49 the diversity and number of visitors to the larger Cytisus flowers. The second, alternative, hypothesis is that the larger Cytisus flowers are more attractive and re- warding than the smaller Genista flowers. Pollina- tors often prefer larger flowers or flowering dis- plays (Campbell 1989; Ohara and Higashi 1994; Schemske and Agren 1995), or larger rewards such as nectar or pollen (e.g., Neiland and Wilcock 1998; Robertson et al. 1999). Flower size can be correlated with nectar production (Brink and de Wet 1980; Harder and Cruzan 1990; Cresswell and Galen 1991), although little is currently known about how flower size correlates with pollen quan- tity or quality among related species. A large-flow- ered species like Cytisus should attract more pol- linators than an otherwise similar, small-flowered species, especially if it offers larger pollen rewards. We measured floral characters and pollen pro- duction in Cytisus and Genista. Armed with infor- mation on the basic floral biology of the two spe- cies, we then tested whether patterns in pollinator visitation, the identity and diversity of visiting in- sect species, and pollinator behavior were consis- tent with the first or second hypothesis. The pur- pose of this study, then, was: 1) to investigate floral traits that may influence attraction, specialization, and pollinator effectiveness; 2) to determine wheth- er visitation rates to these two plant invaders were consistent with predictions based on their floral characters; 3) to determine the identity and origin (native vs. non-native) of insect species pollinating Cytisus and Genista in central California, and 4) to compare the relative “‘quality”’ of pollinators from the introduced range vs. the native range of the in- vaders. METHODS Study Sites We conducted our study at two sites in Marin County, CA. The Mt. Tamalpais site is along the eastern edge of Mount Tamalpais State Park, on an exposed, west-facing slope. This site was dominat- ed by non-native grasses, with patches of Baccharis pilularis, Vinca major, Cotoneaster pannosa, Plan- tago lanceolata, Rubus laciniatus, and Conium ma- culatum. Cytisus and Genista grew interleaved, with Cytisus individuals somewhat more sparse and spread out than Genista. Overall cover of Cytisus and Genista at the site was approximately 15% and 30%, respectively. The China Camp site is located 12 km away from the first site, near China Camp State Park. The site is located in a wide ravine, with mixed evergreen woodland on one side and a steep, NE-facing slope on the other side. Cytisus and Genista grow in patches and as scattered individuals on the ravine bottom and up the NE-facing slope, with a total cover of approximately 15% and 35%, respectively. Other common species included Heteromeles ar- 2002] ia ap , thes IE, Fic. 1. Schematic to illustrate size measurements con- ducted on the flowers of Cytisus and Genista. Flower vol- ume was approximated from a polyhedron using length and width of the keel and height of banner petal. butifolia, Toxicodendron diversilobum, Cortaderia selloana, Baccharis pilularis, Carduus nutans, and non-native grasses. Temperature probes at each site recorded air tem- peratures at 15 min intervals from April 2 through May 31. The two sites were similar for daily max- imum, minimum, and mean termperatures between 9 am and 6 pm. For example, the average daily temperature (CC) was 21.6 in April and 23.1 in May at China Camp, and 21.3 in April and 24.4 in May at Mt. Tamalpais. Floral Characters: Size, Pollen Grain Number, and Accessibility We compared the two species for aspects of their floral biology important to pollinator attraction, re- ward, and utilization. To compare flower sizes, we took three measurements per flower (n = 10 flow- ers for Genista, n = 14 for Cytisus): length of the Keel petal, width of the keel petal, and height of the banner petal. The approximate volume of each flower was calculated as a polyhedron with the ban- ner and keel acting as two planes at right angles, with a third connecting plane (Fig. 1). To quantify pollen production, we collected a single, unopened bud from each of 16 Cytisus (6 Mt. Tamalpais, 10 China Camp) and 19 Genista (7 Mt. Tamalpais, 12 China Camp) plants. We sus- pended and sonicated fully dehisced anthers in a 1% saline solution. We then estimated the number of pollen grains per flower using an Elzone 280-PC electronic particle counter (Micromeritics, Nor- cross, GA). Larger flowers might be more difficult to trip open, limiting access to the flower by smaller pol- linators. To quantify the force required to open the flowers, we hung paper clips sequentially from the keel petals until the flower was tripped open. The paperclips were then weighed and the total mass converted to force. Unpaired t-tests were used to compare the two species for all floral characters. Visitation Rates to Cytisus and Genista Flowers can be tripped only once, after which they remain in an open position. In order to deter- mine the proportion of flowers tripped (an index of Visitation), we marked branches on five individuals PARKER ET AL.: POLLINATION OF CYTISUS AND GENISTA ZT of each species at the two sites. From March 19 through May 21, at one- to two-week intervals, we examined every flower on each branch, recording the number of tripped and untripped flowers. We then marked every open flower with Wite-Out™ to avoid recounting the same flowers at the next cen- sus. Wite-Out@ had no negative effect on fruit pro- duction (Parker and Haubensak 2002). Some un- tripped flowers may have been tripped after being marked and counted (as untripped), leading to an underestimate of the tripping rate. However, we chose this method over the alternative (regular cen- suses without marking flowers), because floral lon- gevity differs between tripped and untripped flow- ers, leading to the possibility that untripped flowers would be double-counted more often than tripped flowers, if not marked. We summed the number of tripped flowers and untripped flowers over the en- tire flowering season (mean flowers per plant = 55 for Cytisus, 153 for Genista), then calculated the total proportion tripped. We used ANOVA to test for differences among plant species and sites, with both species and site treated as fixed effects. For comparison, we plotted the data alongside results from a similar study done in 1997 (Parker and Hau- bensak 2002). Pollinator Observations Throughout the flowering season of 2000 (March 19—May 21), we observed pollinators of Cytisus and Genista at the two sites. The two sites were visited alternately, either on the same day or on consecutive days, and observations were made dur- ing sunny, warm conditions between 9:00 a.m. and 6:00 p.m. In all, we completed 72 hours of obser- vation (36 hours at each site), over 17 days. To maximize our sample size under low visitation rates, we walked haphazard (non-random) transects through the sites, watching and listening for flying insects. We focused on areas where we could see both plant species from the transect. A three-way (pollinator, plant, site) log linear model was used to test for homogeneity of preferences of the two pol- linators (Sokal and Rohlf 1995). For each pollinator, we recorded each attempt to trip a flower, whether that flower was successfully tripped, whether the next flower visited was on the same or a different individual plant, and the dis- tance to the subsequent plant. We calculated the proportion of flowers tripped and the mean number of flowers per plant for each pollinator observed, then averaged these across pollinators for each pol- linator type and each plant species. Only two Bom- bus individuals were observed visiting Cytisus, therefore we could not do a full, two-way analysis using plant species and insect species as factors. Instead, we compared the two plant species for the behavior of Apis only, and compared the two pol- linator species for Genista only. Because the vari- ables (proportion successfully tripped and number DR MADRONO 2000 + - - gin (a) & 1600 - Me S : ) oO iS 1200 + = fo) > 5 800 - | | = | 2 | | Fo 4004 | | 0 a | (b) | © 100000 + | 3 | | é | | = S00004 | | o | 2 ) | Z 60000 ~ ‘3S | 5p | | = 40000 | | | 2 (0) = — 2 Cynsus Genista Fic. 2. Floral traits for Cytisus and Genista: (a) Mean estimated flower volume (n = 14 and 10 for Cytisus and Genista, respectively) and (b) mean number of pollen grains per flower (n = 16 and 19). Error bars represent 1 SE. of flowers tripped per plant) were not normally dis- tributed, we analyzed the data using nonparametric Mann-Whitney U tests. We collected specimens of each pollinator spe- cies and identified them following Thorp et al. (1983). RESULTS Flower Size, Pollen Grain Number, and Accessibility Cytisus flowers were significantly larger than Genista flowers; length of the keel petal (mm + 1 SD) was 18.8 + 1.4 compared to 6.7 + 0.5 for Genista (df = 22, t = 27.1, P < 0.0001). The es- timated volume of Cytisus flowers (1450 mm? + 364 SD) was an order of magnitude larger than that 0.035 + — 0.030 - ] 00254 | 0.020 - 0.015 - 0.010 5 | | 0.000 -—1— 0.005 + Force required to trip flower (Newtons) Genista Cyrsus Fic. 3. Force required to access (trip open) flowers of Cytisus and Genista, determined with a mechanical assay. Error bars represent 1 SE. [Vol. 49 IO = 5 oa VISUS Bo Genista a) 2000 08 4 é 06 5 ws) OA 4 oO a. | = = O24 o 3 © rm 00 az 4 fo) & b) 1997 & (2 = a a. 2 Ou 06 5 | e | = O04 - 02 4 : | 09 +1 . China Camp Mt. Tamalpais Fic. 4. Proportion of flowers tripped for Cytisus (open bars) and Genista (filled bars) in two populations. a) Data from 2000, collected from 5 individuals per population; b) Data from 1997, collected from 40—60 individuals per population. Data from 2000 redrawn from Parker and Haubensak (2002). Error bars represent 1 SE. for Genista-(i32 = 27; Bigs 2:'dti = 2207 — isles P < 0.0001). Cytisus produced more than four times as many pollen grains per flower as Genista (Fig. 2, df = 33, t = 8.0, P < 0.0001). In the mechanical assay, the two species required significantly different amounts of force to trip the flowers (df = 56, t = 2.80, P = 0.007). We used more than twice as much force in opening flowers of Cytisus (0.026 N + 0.005 SD) as in opening flowers of Genista (0.013 + 0.006) (Fig. 3). Visitation Rates to Cytisus and Genista At both sites, large-flowered Cytisus had a sig- nificantly higher proportion of tripped flowers than did small-flowered Genista (Fig. 4a, Table 1). These results from the 2000 season confirmed the same pattern seen in 1997 (Fig. 4b, Parker and Haubensak 2002). Therefore, the greater pollination success of Cytisus seems to be generalizable over both space and time. There was no significant main effect of site (Table 1). Rather, for Cytisus, more TABLE 1. ANALYSIS OF WARIANCE OF PROPORTION OF FLOWERS TRIPPED (I.E., WISITED) OVER THE ENTIRE SEASON FOR CYTISUS AND GENISTA AT TwoO SITES. Source df SS F Pp Species 1 0.80 61.1 <0.0001 Site 1 0.01 0.53 0.48 Species X Site 1 0.07 5.07 0.039 Residual 16 0.21 2002] TABLE 2. NUMBERS OF POLLINATORS OBSERVED IN 72 HOURS OF OBSERVATION AT TwoO SITES ON THE MARIN PEN- INSULA. Observations were done along haphazard transects through the sites, in areas where both plant species were within visual and auditory range. Mt. Tamalpais China Camp Insect species Cytisus Genista Cytisus Genista Apis mellifera 19 3} q. 12 Bombus vosnesenskii 1 i>) 1 0 Xylocopa californica 0) 1 0 0 Total 20 19 8 12 flowers were tripped at China Camp, while for Gen- ista, more flowers were tripped at Mt. Tamalpais (Fig. 4a), resulting in a significant population xX species interaction effect (Table 1). Pollinator Observations In 72 hours of observation at the two sites, three bee species were seen pollinating Cytisus and Gen- ista: Apis mellifera, Bombus vosnesenskii, and Xy- locopa californica (Table 2). We observed a total of 59 pollinators, which made 342 effective visits to flowers (in which the flower was tripped). Given that hundreds to thousands of flowers were being watched at any one moment, this represents a very low pollination rate in terms of visits/flower/hour. Apis was the numerically dominant pollinator ob- served at both sites, although Genista at Mt. Tam- alpais was visited most by Bombus (Table 2). Xy- locopa was observed only once and is left out of subsequent analyses. Results of the three-way log linear model indicated that bee species preferences for the two broom species differed across the two sites (G[Williams] = 8.18, df = 3, P < 0.005). Therefore, we examined these preferences for each site separately. At Mt. Tamalpais, Apis primarily visited Cytisus, while Bombus primarily visited Genista (G[Williams] = 26.48, df = 1, P < 0.001). At China Camp, there were too few Bombus visits to include in the analysis, but a chi-square test in- dicated that Apis had no significant preference for Cytisus or Genista (chi-square = 1.32, df = 1, P > 0.10). Therefore, we did not find evidence to sup- port the prediction that the small-flowered Genista was accessible to a greater number of pollinator species. Nor did we find that the larger pollinators (Bombus) tended to visit the larger flowers (Cyti- SUS). Before flying to a new individual, bees (all spe- cies combined) visited on average 6.6 (+10.6) flowers/plant of Cytisus and 3.9 (+3.8) flowers/ plant of Genista, a difference that was not statisti- cally significant (Z = 0.25, P = 0.80). When con- sidered alone, Apis did not differ in its pattern of movement on the two plant species (Fig. 5, Z = 0.22, P = 0.83). Although Apis tended to visit more flowers per plant than Bombus (Fig. 5), this differ- PARKER ET AL.: POLLINATION OF CYTISUS AND GENISTA 29 10 [|] Cynisus 8 i Genista Number of flowers tripped per plant Bombus Apis Fic. 5. Number of flowers successfully visited (tripped) per individual plant before flying to a new plant, for Bom- bus observed on Cytisus (n = 2) and Genista (n = 15) and Apis observed on Cytisus (n = 22) and Genista (n = 15). Error bars represent 1 SE. ence was not Statistically significant (Genista only, Z = 1.20, P = 0.23). We used Apis visitation success (ratio of flowers tripped to flowers attempted) as a measure of the accessibility of the two flower types; pollinators were more successful at accessing flowers of the small-flowered Genista than the large-flowered Cy- tisus (Fig. 6, Apis only, Z = 2.13, P = 0.03). The proportion of Genista flowers successfully tripped did not differ for the two pollinator species (Fig. 6, Z = 1.0, P = 0.32). The proportion of Cytisus flow- ers successfully tripped appeared to be higher for Bombus than for Apis (Fig. 6). This difference was statistically significant (Z = 2.09, P = 0.03), but the small sample size for Bombus (N = 2) suggests caution in interpreting this result. DISCUSSION Do Floral Characters Predict Which Invasive Plants Should Be Most Attractive to Pollinators and Which Should Be Most Pollen Limited? We started with two alternative hypotheses for how floral traits could affect pollinator attraction and visitation in this system. The first hypothesis (—— 2 ios —— = _— | Cytisus Bi Cenista Prop. of flowers successfully tripped oO Oo Bombus Apis Fic. 6. The mean proportion of flower visits attempted in which the flower was successfully tripped open, for individual Bombus and Apis pollinators observed at flow- ers of Cytisus (open bars) or Genista (filled bars). Each pollinator observed could involve visits to flowers on one plant or multiple plants. Error bars represent 1 SE. 30 predicted that because of the unusual “‘tripping”’ mechanism in these two species, the larger Cytisus flowers could be more difficult for pollinators to handle, which would reduce the number of insect species capable of visitation. Cytisus would then be relatively specialized on larger insects, and polli- nator limitation should be more prevalent in Cytisus than in Genista (assuming an equal and variable pool of pollinator species for the two plants). In fact, we did find evidence that Cytisus flowers are more difficult to access than Genista flowers, both from our mechanical assay and from observations of Apis visitation. However, this difference did not result in a difference in the pollinator assemblage for the two plants. We did not find that small soli- tary bees or flies visited Genista, although they are common in the area (G. Lebuhn, personal com- munication), and we did not find that larger bees specialized on Cytisus. Rather, while Apis com- monly visited both plant species, the larger Bombus and Xylocopa visited primarily Genista. Recent work in Great Britain (Stout 2000) found that very large Bombus individuals (>20 mm) were less ad- ept at accessing Cytisus flowers than were smaller Bombus. This suggests, in concert with our results, that the largest bees may have trouble handling Cy- tisus flowers and may in fact prefer Genista flowers While we did not find that the differences in flo- ral accessibility between the two invaders translated to an effect on pollinator composition in Marin County, this effect could be important in other sys- tems. For example, one of us (Parker 1997) ob- served that Cytisus was rarely successfully tripped by Apis in Washington State. Temperature can af- fect floral accessibility: Cytisus flowers are more difficult to trip in cold compared to warm temper- atures in the lab (B. Burley, R. Martin, and K. M. Karoly unpublished data). The Washington research suggests that in colder climates, the difficulty of tripping Cytisus flowers could contribute to pollen limitation. Our second hypothesis postulated that pollinator visitation would be based on floral attractiveness. The prediction was that the larger flowers and greater pollen rewards of Cytisus could increase its attractiveness to pollinators, increasing its success in competing with resident plants for pollinator ser- vices relative to Genista. We found, both in 1997 and 2000, that visitation rates to the two plant spe- cies were consistent with this hypothesis. Cytisus flowers were more frequently tripped in both years and at both study sites. Given that Genista tends to produce more flowers than Cytisus on plants of the same size (Parker and Haubensak unpublished data), pollinators in this study appeared to be more attracted by the rewards of single flowers than by the size of the overall display. Genista might have ways of compensating for its lower pollinator visitation. Small flowers some- times represent reduced allocation to attraction in an evolved syndrome including increased selfing MADRONO [Vol. 49 and autogamy and lower inbreeding depression (Piper et al. 1986; Rathcke and Real 1993; Parker et al. 1995; Brunet and Eckert 1998). Genista does not exhibit high levels of autogamy (Parker and Haubensak 2002). However, compared to Cyfisus, it does show less inbreeding depression at seed-set, consistent with the syndrome of increased selfing and decreased allocation to attractive structures (Parker and Haubensak 2002). Thus when pollina- tors tend to visit multiple flowers on a plant, Gen- ista may be able to take advantage of those visits more effectively than Cyfisus. Are Plants Better “‘Matched”’ to Pollinators from Their Native Range? If the floral traits of plants have evolved in re- sponse to pollinators in their native range, one might expect to see that a pollinator introduced from the same region as an introduced plant would be a higher quality pollinator, or better match, for that plant. We found no evidence to support this prediction in our study. Apis mellifera, native to Europe, was not better at tripping flowers than the locally native Bombus vosnesenskii. We did not fol- low the fate of each tripped flower to determine relative pollen transfer and fruit set for the different pollinators (see Schemske and Horvitz 1984). How- ever, the European Apis and native Bombus did not differ significantly in their tendency to promote out- crossing by moving between plants instead of with- in plants. Given that most plant-pollinator interactions are thought to have arisen from diffuse coevolution of guilds rather than coevolution between pairs of spe- cies (Jordano 1987; Pellmyr 1992), these findings are not particularly surprising. European species of Bombus probably occurred throughout the evolu- tionary history of these broom species, and may have influenced the evolution of their floral mor- phology, and North American and European Bom- bus are probably similar in terms of their pollina- tion value to broom. Although we lack extensive information about the degree to which Cytisus and Genista are pollinated by Apis in their native range, one recent study in Great Britain recorded that flowers of Cytisus there were tripped primarily by Bombus, while Apis only visited previously tripped flowers (Stout 2000). More comparative studies of | plant-pollinator interactions in native vs. introduced ranges are needed to better understand the role of these mutualisms in the invasion process. We are aware of only one case in which a highly specific pollinator was left behind during invasion (e.g., fig wasp invasion in Florida, Nadel et al. 1992). Invasional Meltdown? Both Cytisus and Genista can be pollen limited (Parker 1997; Parker and Haubensak 2002); there- fore, attracting pollinators plays a critical role in | assuring reproduction. We found that both native ——SSSEeeee SSS 2002] and non-native insects pollinate these invaders. Apis mellifera, native to Europe, was by far the dominant visitor at the China Camp site, and was also more common than Bombus at Mt. Tamalpais. It has recently been suggested that positive inter- actions among invaders may commonly facilitate new invasions or exacerbate the spread or impacts of established invaders, leading to non-linear “‘in- vasional meltdown’? (Simberloff and Von Holle 1999). In our system, an introduced pollinator ap- pears to be facilitating the invasion of two intro- duced shrubs. : Recent concern over the environmental impacts of Apis introduction has focused on the fates of native pollinators (Roubik 1982; Buchmann and Nabhan 1996; Goodell 2000). We point to another potential conservation issue, the effect of Apis in- troduction on the spread of noxious weeds in nat- ural ecosystems. In keeping with this warning, re- cent work by Barthell et al. (2001) found that pol- lination by Apis contributed up to half of the seed produced by the noxious invader Centaurea solsti- tialis (yellow star thistle) in California. Beekeeping is common at the suburban-rural interface. The bees at our sites may have come from feral colonies, or beekeepers may have been responsible for domestic colonies in nearby residential or agricultural areas. 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Our analyses revealed a lack of interregional divergence of stand characteristics between the Klamath Mountains and southern Sierra Nevada despite subspecific designa- tion and recently identified genetic divergence. Bray-Curtis ordination and hierarchical cluster analyses identified four stand types—1) stands dominated by foxtail pine, 2) stands with foxtail pine and whitebark pine, 3) stands with foxtail pine and red fir, and 4) mixed stands with foxtail pine, red fir, and western white pine. Sub-regions within the Klamath Mountains differed in foxtail pine relative density, conifer diversity, and substrate heterogeneity. Further analyses of the Klamath Mountains stands identified an inverse relationship between foxtail pine importance and conifer diversity. Interactions between the moun- tain island effect and substrate heterogeneity were inferred as regulatory mechanisms for foxtail pine importance and conifer diversity, but further research is needed to determine causal relationships from our correlations. Key words: foxtail pine, conifer diversity, mountain island effect, substrate heterogeneity, Klamath Moun- tains Foxtail pine (Pinus balfouriana Grev. & Balf.) is a California endemic conifer found in two isolated areas of the state separated by 500 kilometers. In extreme northern California it grows in the central ranges of the Klamath Mountains, while the south- erm population is centered on the Cottonwood Basin and surrounding areas in the southern Sierra Ne- vada. The Scottish botanist and explorer John Jef- fery discovered it in 1852 on a botanical recon- naissance of the Klamath Mountains. Professor John H. Balfour and Dr. R. K. Greville subsequent- ly prepared the original description and illustrations for the Oregon Association (Mastrogiuseppe 1972). Since its discovery, however, there has been a pau- city of ecological research into the biology of this subalpine tree. Ball (1976) highlights the need for research into the factors determining the ecological patterns of foxtail pine. Foxtail pine is a five-needle haploxylon pine placed within the subsection Balfourianae Engelm. along with Great Basin bristlecone pine (Pinus lon- gavea D. Bailey) in the western Great Basin and the Rocky Mountain bristlecone pine (Pinus aris- tata Engelm.) in the eastern Great Basin and Rocky Mountains. Mastrogiuseppe (1972) argued that two allopatric subspecies of foxtail pine exist upon dif- ferences in mean cone length, seed wing length, cotyledon number, and needle resin duct spacing. Mastrogiuseppe and Mastrogiuseppe (1980) subse- ' Present address: Department of Botany, Box 355325, University of Washington, Seattle, WA 98195. quently named the southern population Pinus bal- fouriana ssp. austrina. An 11 loci allozyme analysis of stands within the northern population revealed high levels of genetic differentiation among stands (Oline et al. 2000). This genetic diversity was significantly greater than the differentiation between the northern and south- ern populations and was hypothesized to be a func- tion of the mountain island effect, genetic drift, possible serpentine soil adaptation, and high conifer richness (Hamrick et al. 1994; Oline et al. 2000). Foxtail pine and Great Basin bristlecone pine from the White Mountains exhibit strong anatomi- cal and morphological similarities suggesting hy- bridization (Mirov 1967; Bailey 1970). Successful experimental crosses support this proposed hybrid- ization based on morphological continuity (Critch- field 1977). Numerous hypotheses concerning the origin of the subsection Balfourianae, foxtail pine, and its subspecies have been offered (Mastrogiu- seppe 1972; Critchfield 1977; Raven and Axelrod 1978). Most hypotheses, however, date the initial divergence of the bristlecone/foxtail pine ancestor to the Oligocene and the disjunction of foxtail pine within California to the Xerothermic period of the Holocene approximately 8000 years ago (Critch- field 1977; Raven and Axelrod 1978) or the Sierran orogeny approximately 2 million years ago (Mirov 1967; Bailey 1970). Population and community-level research on foxtail pine is sparse. This pine is intolerant of shade at all stages of growth, an inhabitant of var- ious high elevation substrates, and is a major to minor component of subalpine forests and wood- 34 MADRONO lands in both the Klamath Mountains and southern Sierra Nevada (Ryerson 1983; Sawyer and Thorn- burgh 1988). Extreme old age is achieved on xeric high altitude locations and trees can reach a maxi- mum age of approximately 2000 years in the south- ern population (Ball 1976; Scuderi 1987). Trees from the northern population, however, only attain maximum ages of approximately 800 to 1,000 years as a result of widespread heart rot and periodic fire (Mastrogiuseppe 1972). Foxtail pine typically occurs in low diversity stands with whitebark pine (Pinus albicaulis En- gelm.), lodgepole pine (Pinus contorta Loudon ssp. murrayana Critchf.), western white pine (Pinus monticola Douglas), red fir (Abies magnifica Andr. Murray), and mountain hemlock (Tsuga mertensi- ana Carriere) (Mastrogiuseppe 1972; Ryerson 1983). Common ground layer associates include dry site graminoids, herbs, and shrubs such as Agrostis, Arabis, Arcotstaphylos, Carex, Ceano- thus, Eriogonum, Festuca, Juncus, and Sedum (Ryerson 1983; Sawyer and Keeler-Wolf 1995). Foxtail pine in the Klamath Mountains forms ex- tensive stands at high elevations on isolated peaks and ridges with mafic and ultramafic substrates where it is the dominant tree. Exceptions occur in the China Peak and Russian Peak areas where it is associated with as many as six other conifer species in a mixed subalpine forest type (Sawyer and Kee- ler-Wolf 1995). Conifers such as Jeffrey pine (Pi- nus jeffreyi Grev. & Balf.), incense-cedar (Caloced- rus decurrens (Torrey) Florin), and Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) form a minor component in these stands. In the southern Sierra Nevada, foxtail pine forms extensive low di- versity stands at high elevations on granitic and metamorphic substrates. Common associates in- clude whitebark pine, limber pine (Pinus flexilis James), red fir, and western white pine as described by Vankat (1970) in Sequoia National Park as a foxtail pine forest type. Limited research suggests that foxtail pine’s dis- tribution is determined by variation in substrate, cli- mate, and interspecific interactions with other co- nifers (Mastrogiuseppe 1972; Ryerson 1983; Scu- deri 1987; Oline et al. 2000). The low competitive ability of foxtail pine and the high regional conifer species richness of the Klamath Mountains may il- lustrate restriction of foxtail pine to safe sites on ultramafic substrates and granitic boulder fields where shade tolerant conifers cannot shade out in- tolerant foxtail pine trees (Ryerson 1983; Sawyer and Thornburgh 1988). Previous research has identified examples of eco- logical differences between the two populations. Foxtail pine stands in the southern Sierra Nevada achieve their highest densities on northern slopes (Ryerson 1983), which contradicts the typical southern and eastern slope success observed in the Klamath Mountains (Mastrogiuseppe 1972). Sur- prisingly the range of foxtail pine, commonly de- [Vol. 49 scribed as a Pleistocene relict, has been document- ed as expanding to lower elevations in the southern Sierra Nevada (Ryerson 1983). This expansion may be a function of climate change, low conifer rich- ness, demographic and environmental stochasticity, or a combination of these processes (Mastrogiusep- pe 1972; Ryerson 1983; Scuderi 1987). We documented environmental and composition- al gradients related to the level of dominance of foxtail pine among stands in the Klamath Moun- tains. We also compared these stands to those of the southern Sierra Nevada using the studies of Vankat (1970) and Ryerson (1983). These compar- isons allowed us to analyze the disjunct foxtail pine populations at several levels and to make inferences about the role of interspecific interactions, geologic substrate, and climate in the distribution and dom- inance of foxtail pine at the intraregional and local levels in the Klamath Mountains. METHODS Study areas. The Klamath Mountains are com- posed of a complex set of predominantly southwest to northeast trending mountain ridges separated by deep canyons and valleys that encompass approxi- mately 30,300 km? of northern California (Fig. 1). The main mountain ranges, from south to north, are the South Yolla Bolly Mountains, North Yolla Bol- ly Mountains, Trinity Alps, Salmon Mountains, Scott Mountains, Trinity Mountains, Marble Moun- tains, and Siskiyou Mountains. Summits average from 1500 to 2100 m in elevation, with a maximum of 2750 m at Mount Eddy. The major watersheds include the Sacramento, Trinity, Salmon, Scott, Klamath, and Smith Rivers. Climate patterns con- form to a modified mediterranean climate type with long, wet winters and generally dry summers. Ay- erage precipitation ranges from 125 cm to 175 cm, with thunderstorms and lightning developing in late August and early September (Major 1988). Plant assemblages range from low elevation chaparral, woodlands and forests, to patches of alpine plants, with a number of endemic and relict populations of vascular plants on ultramafic substrates (Walker 1954; Kruckeberg 1992). The southern Sierra Nevada is a linear north- west-southeast trending mountain range bounded on the west by the San Joaquin Valley and on the east by the White Mountains and the Great Basin (Fig. 1). Over 285 peaks reach elevations of 3600 m, over 140 exceed 3900 m, while 11 peaks top 4200 m (Kruse 1990). Major watersheds include the many forks of the Kern, Kings and San Joa- quin Rivers. Regional climates are influenced by local topography, but all resemble a modified Mediterranean climate with moderately wet win- ters and dry summers broken by periods of after- noon thunderstorms. Annual precipitation ranges from 35 cm to 115 cm on the western crest and from 50 cm to 75 cm on the eastern crest (Major ) . | . | | | | | 2002] 32 pha GMM Pinus balfouriana ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 35) Ee eines i x $32 : Qo x isolated occurrence | o “_ 2 Limits of Pras aristetet aS ow -a | = 9 Igo 200 300 MILES oe - - se a . vad OO 260 $00 400 KILOMETERS | | We jams t = 7 720 ne 1S ; Fic. 1. Distribution of foxtail pine in the Klamath Mountains and Sierra Nevada by Griffin and Critchfield (1972). MM = Marble Mountains, RP = Russian Peak, TA = Trinity Alps, TM = Trinity Mountains, YB = Yolla Bolly Mountains. 1988). Plant assemblages range from low eleva- _ tion pinyon pine woodlands to alpine tundra on the eastside and from foothill woodlands, mon- tane and subalpine forests, to alpine vegetation _ types on the westside. Sampling techniques. We identified five sub-re- gions where foxtail pine was dominant within the Klamath Mountains—the Yolla Bolly Mountains, the Trinity Alps, the Trinity Mountains, Russian Peak, and the Marble Mountains (Table 1). Each 26 MADRONO TABLE 1. SUB-REGIONAL LOCALITIES OF 15 FOXTAIL PINE STANDS IN THE KLAMATH MOUNTAINS. SUB-REGIONS AND LOCALITIES WERE BROADLY DEFINED AND DO NOT HAVE DISs- TINCT BOUNDARIES. Sub-region Locality Stands Marble Mountains Lake Mountain 2 Russian Peak Russian Peak 1 Trinity Alps Granite Peak 2 Seven-Up Peak 1 Union Creek D Trinity Mountains Mt. Eddy 2 China Mountain/ Crater Lake 2 Yolla Bolly Mountains North South—Mt. Lynn 1 sub-region contained distinct localities where fox- tail pine density was sufficiently high for sampling. At each locality we randomly identified stands pri- marily through homogeneity of species composi- tion and secondarily through homogeneity of geo- morphology. Stand areas ranged from 8 ha to 28 ha. A total of 15 stands were sampled from the 10 localities in the 5 sub-regions from May 22 through September 16, 2000. We used the point centered quarter (PCQ) meth- od to sample each of the 15 stands in order to es- timate tree density, basal area, frequency, and im- portance values by species. This method is a plot- less sampling technique based upon the placement of random points along pre-determined transects perpendicular to contour lines within each stand. Trees were defined as single conifer individuals with diameter measured at breast height for trees taller than 1.37 m, at ground level for trees below 1.37 m in height, and as an average for trees with multiple trunks. Ryerson (1983) sampled 15 marginal foxtail pine stands located in the southern Sierra Nevada using the PCQ method in a similar manner to estimate conifer importance values. Vankat (1970) conduct- ed linear transect sampling of foxtail pine forest type within Sequoia National Park to estimate stand and species-specific tree density, basal area, and frequency. To determine the number of points needed to ad- equately estimate stand density, we conducted pre- sampling of two stands in the North Yolla Bolly Mountains following Bonham’s (1989) method. We determined that 22 points were adequate to estimate stand-level parameters with an error of 10%. We increased this number to 25 to ensure reliable es- timation, but were only able to establish 15 points in the Marble Mountains stands due to their small Size. To generate estimates of tree density, basal area, and frequency, we placed 25 points along four to six transects that were evenly spaced throughout the stand. Spacing among points was dependent upon stand area. We followed Bonham’s (1989) [Vol. 49 protocol at each point, yielding 100 sampled trees per stand. The diameter at breast height (DBH) was measured for each tree, and the average diameter and stem number were obtained for trees with mul- tiple trunks. We measured percent slope, aspect, el- evation, and identified bedrock type. Cover of boul- ders (>50 cm), cobbles (10 cm—S50 cm), gravels (<10 cm), and organic material was estimated us- ing the Braun-Blanquet scale (Bonham 1989). Estimators and statistical analyses. The PCQ method allowed unbiased estimation of tree density and basal area within each stand (Pollard 1971). From these values we calculated stand and species- specific tree density, basal area, frequency, and im- portance values (Kent and Coker 1992). Importance values are composite estimates based upon the summation of relative estimates of species density, frequency, and basal area (Bonham 1989). These values could therefore range from a minimum of zero, if the tree species was absent, to a maximum of 300 if it was the only species present. We placed foxtail pine trees into ten size classes based upon diameter measurements. Using a simple linear regression model constructed from tree ring counts obtained from Mastrogiuseppe (1972), we related logarithmically transformed values of tree diameter to age by the following equation: Log(age) = 1.1571(Log[DBH]) + 0.3762 (1) This model was significant (ANOVA: F,4,; = 277.0459, P = 0.000001) and had a high degree of explanatory power (R* = 0.8738). Hundred-year age classes could be correlated to size classes with confidence and reproductive success inferred from relative density of the first age class (Ryerson 1983). Survival among age classes was calculated as the ratio of the density of age class x to the density of age class x + 1. We used the Shannon-Wiener Index (10g,)) to measure substrate heterogeneity and tree diversity using median species-specific relative density val- ues for conifer diversity. We computed Jaccard sim- ilarity coefficients, weighted by species-specific im- portance values, among all stands to obtain average within and among sub-regional stand similarities within the Klamath Mountains. We subjected the data to a variety of statistical analyses. Significance levels for hypothesis tests were set at a = 0.05 or 0.10. A modified t-test checked for significant dif- ferences among index values (Zar 1999). Two-sam- pled t-tests and Mann-Whitney U tests established interregional differences and similarities for stand characteristics using Ryerson’s (1983) and Vankat’s (1970) data sets. Variation around the mean of these estimates, if the data were normally distributed, was compared between regional populations using Har- tley’s equal variance test. We used simple linear regression analyses and General Linear Models (GLM) ANOVA’s to identify trends within and dif- ferences among stand characteristics and environ- mental variables by sub-regions within the Klamath 2002] ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 37 TABLE 2. SUMMARY OF THE ENVIRONMENTAL VARIABLES AND STAND CHARACTERISTICS OF 15 FOXTAIL PINE STANDS FROM THE KLAMATH MOUNTAINS BY SUB-REGIONS. Conifer diversity and substrate heterogeneity are reported as Shannon- Wiener Index (log,, base) values. Density given in trees/hectare and basal area in m*/hectare. Basal Area Elevation Slope Substrate Conifer Conifer Stand Density area (ha) (m) Aspect (%) heterogeneity diversity richness Yolla Bolly Mountains 1 238 45 8.6 2365 WSW 28.0 0.09 0.39 3) 2, Sil 10 25.4 2260 WSW BOF] 0.38 0.44 5 3 85 14 L523 2335 NNW 42.6 0.18 Oi 3 Trinity Alps 4 nO DES 10.1 2240 WSW 29.0 0.29 0.42 5 5) Di. Oa 11.4 2345 WNW 39:6 0.29 0.44 5 6 133 36 28.0 DUES WNW 58).3) 0.29 0.26 3 gl eZ Dey 11.2 2202 WNW 40.1 0.25 0.42 3 8 190 38 28.0 2004 NNE 153 0.41 0.52 6 Trinity Mountains 9 252 44 47.5 2358 WNW 25.8 O33 OZ 6 10 207 8 24.5 2376 SSE 32) 0.41 0.61 6 iN 92 26 28.1 2500 WNW S57 0.41 0.42 5 12 381 I 14.0 2205 ESE Te 0.41 0.69 6 Russian Peak 13 315) 24 12.8 2180 NNW S85 0.44 0.84 8 Marble Mountains 14 all 25 8.2 2003 SSE BES) 0.41 0.66 6 15 198 10 Sal 2033 NNE 35.0 0.20 0.50 6 Mountains. Average within and among sub-region- al Jaccard similarity coefficients were compared us- ing GLM ANOVA with Fischer’s LSD grouping algorithm among sub-regions with more than two stands. We used hierarchical cluster and Bray-Curtis or- dination analyses to reduce dimensionality within the data sets and to analyze multivariate correla- tions among stand compositional and environmen- tal variables (Kent and Coker 1992). Hierarchical cluster analyses provided dendrograms based on Euclidean distance, scaled by standard deviations, using species importance values. Clusters were identified with the group average (unweighted pair- groups) algorithm. We employed Bray-Curtis ordi- nation analyses to evaluate the results of the hier- archical cluster analyses and to correlate stand groupings with environmental variables using the Sorenson similarity coefficient based upon species importance values. Biplots were overlain with com- positional and environmental variable vectors and correlations with the first two axes were computed (McCune 1993). RESULTS Summary. Approximately 1,420 trees were sam- pled in the determination of importance values for 11 conifer species sampled within 15 stands located in the Klamath Mountains. Stand densities ranged from 50 trees/ha in the North Yolla Bolly Moun- tains to 381 trees/ha at China Mountain (Table 2). Stand basal area ranged from a minimum of 8 m?/ ha at China Mountain to a maximum of 45 m?/ha at Mt. Lynn (Table 2). Foxtail pine was the most important conifer in 14 out of the 15 stands with a regional average importance value of 168, a mini- mum of 46 at Crater Lake, and a maximum of 241 at Seven-Up Peak (Table 3). The first age class (<100 years) dominated (=50% relative density) the age class distribution of foxtail pine in 9 out 15 stands. Maximum tree age approached 1000 years for a few foxtail pine trees scattered throughout stands. Average stand abundances by age class within the Klamath Mountains assumed J-shaped distributions and differed (ANOVA: F,14;; = 120.06, P = 0.000001). This pattern was also ob- served in the southern Sierra Nevada (Ryerson 1983). Klamath Mountains. The common tree species were foxtail pine, red fir, and western white pine with importance value dependent upon geographic sub-region. Foxtail pine achieved higher densities on steep, high elevation slopes with western aspects (Tables 2, 3). Whitebark pine was primarily found in stands on high elevation peaks with granitic or glacial till substrates. Mountain hemlock tended to be most important on northern slopes at lower el- evations and east slopes at higher elevations. Co- nifers typical of lower elevations, such as incense- cedar, Jeffrey pine, and Douglas-fir, were found in stands with elevations below 2200 m in elevation or western and southern aspects. Conifer diversity was variable throughout the re- gion, but was greatest in northern-most stands (Ta- ble 2). Stands in the Yolla Bolly Mountains and southern Trinity Alps had lower diversity values, while stands in the Trinity and Marble Mountains had higher diversity values. Diversity values ranged 38 MADRONO [Vol. 49 TABLE 3. CHARACTERISTICS OF FOXTAIL PINE WITHIN 15 STANDS FROM THE KLAMATH MOUNTAINS BY SUB-REGIONS. Density given in trees/hectare, basal area in m7/hectare, and relative values in percent. Relative Stand Density density Yolla Bolly Mountains 1 ISS 65 D 52 63 3 62 73 Trinity Alps 4 98 Syil 5 2 47 6 109 82 d 53 47 8 116 61 Trinity Mountains 9 153 61 10 79 38 11 13) 59 12 69 18 Russian Peak 13 69 2D Marble Mountains 14 74 43 15 109 5) from 0.25 at Seven-Up Peak to 0.83 at Russian Peak. These two values were statistically different (t; = 43.9796, P < 0.00001) according the t-test proposed by Hutcheson (1970). Conifer richness was as low as 3 species in the Trinity Alps stands and as high as 8 species in the Russian Peak stand. Foxtail pine inhabited slopes with a variety of aspects. A large proportion of stands had northwest and southwest aspects with foxtail pine importance higher on southwest facing slopes (Tables 2, 3). Substrate heterogeneity measured with the Shan- non-Wiener Diversity Index ranged from a low of 0.09 in the North Yolla Bolly Mountains to 0.44 at Russian Peak (Table 2). Inceptisols developed from mafic and ultramafic geologic substrates were the primary soil type within stands. Pockets of granitic and metasedimentary rocks were prominent in the Yolla Bolly Mountains stands and at Russian Peak. Evidence of glaciation was present in high eleva- tion stands with glacial till as an important substrate at stands in the Trinity Alps, Russian Peak, and Trinity Mountains. Elevations ranged from a low of 2000 m at Lake Mountain to a high of 2500 m at China Peak with a regional average of 2247 m (Ta- blew): Simple linear regression analyses identified nu- merous compositional and environmental gradients among stands located in the Klamath Mountains (Fig. 2). A south to north latitudinal gradient was identified as significantly influencing foxtail pine relative density, conifer diversity, species richness, substrate heterogeneity, and foxtail pine survival to the second age class (F,,4 > 4.35, P < 0.05, R* > 0.30). Conifer diversity negatively influenced fox- tail pine importance (Fig. 2A). Substrate heteroge- neity, in turn, was identified as positively influenc- Basal Relative Relative Importance area frequency dominance value 38 46 95 207 7 48 89 199 10 56 90 220 ts) 48 Syl 156 13 44 51 142 31 61 98 241 20 46 80 7/3) 33 44 81 187 38 45 96 202 5) 55 68 139 21 46 88 195 3) 18 10 46 9 26 38 85 7 31 89 163 4 40 64 IS) ing conifer diversity (Fig. 2B). This relationship was dependent upon Pleistocene glaciation with stands on glaciated surfaces exhibiting a stronger correlation with high conifer diversity (glaciated R? = 0.6181, unglaciated R? = 0.3002). GLM ANOVA’s of compositional and environ- mental variables revealed sub-regional differentia- tion. Elevation, conifer species richness and diver- sity, whitebark pine relative density, and foxtail pine survival to the second age class significantly differed among the five sub-regions (ANOVA: F,,, > 3.25, P < 0.05). In general, foxtail pine survival was highest in stands with lower conifer diversity. Stands were more homogeneous, as measured with average sub-regional Jaccard similarity coef- ficients, within sub-regions versus among sub-re- gions (Fig. 3). In general, within sub-region Jaccard similarity coefficients were greater than among sub- region coefficients with the Marble Mountains be- ing similar to all other sub-regions (ANOVA: F3 157 = 17.50, P = 0.0000009). Russian Peak was most similar to the Trinity Mountains and Marble Moun- tains, while least similar to the Trinity Alps and Yolla Bolly Mountains. Similarity was not corre- lated to distances among sub-regions, however the extreme northern and southern stands were among the least similar. The recently disturbed stand (e.g., Crater Lake, stand 12) was the least similar to oth- ers at both within and among sub-regional levels. The Bray-Curtis ordination of the Klamath stands differentiated stands with high conifer rich- ness from those with high foxtail pine importance values (Fig. 4). Axis I in the ordination was highly and variously correlated with mountain hemlock (R = —(0.837), western white pine (R = 0.564) and foxtail pine (R = 0.547) importance values. Axis II mG 2. _ 0.6399). Stands with serpentine soils are represented by filled points. (B) Relationship between substrate heterogeneity _ and conifer diversity grouped by glaciation history (dashed line = 2002] (A) 250 5 225 - 200 - “< oO e Foxtail pine importance N (Sy) 205 (0) - 7s T T ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 38) T U T T 0.35 0.40 T T 0.20 0.25 0.3 I T 0.45 0.50 055 060 065 070 0.75 0.80 0.85 0.90 Conifer Diversity ‘= fo) 1 Conifer Diversity —) (3)] | 0.4 5 oO Glaciated m Unglaciated - R*=06181 T 0.20 0.25 0.30 0.35 0.50 Substrate Heterogeneity 0.15 0.40 0.45 (A) Relationship between conifer diversity and foxtail pine importance (F, ,, = 23.096, P = 0.000343, R? = unglaciated regression, solid line = glaciated regression). See text for definitions of conifer diversity and substrate heterogeneity. was similarly correlated with western pine (R = —0.617), whitebark pine (R = —0.616), and foxtail pine (R = 0.603) importance values. These two axes had relatively high explanatory power (R? = 0.739). Stands tended to be broadly grouped by sub-region and composition (Fig. 4). Crater Lake was unique with extremely low foxtail pine impor- _ tance and high conifer species richness. This stand was selectively logged and burned in the last half century. Southern Sierra Nevada. Using Ryerson’s (1983) data set, stand density ranged from a low of 72 trees/ha to a high of 881 trees/ha, while stand basal area ranged from a low of 9 m*/ha to 646 m*/ha (Table 4). Extremely high density and basal area values were associated with whitebark pine domi- nance and may have been an artifact of sampling trees with numerous trunks. A wide range of im- portance values for foxtail pine, low species rich- ness, and high elevations characterized these stands (Table 4). Using Vankat’s (1970) data set, stand density ranged from a low of 200 trees/ha to a high of 700 trees/ha, while stand basal area ranged from 15 m7/ha 40 | MADRONO [Vol. 49 97.5 95.0 92.5 90.0 - 87.5 85.0 Average Jaccard Similarity (%) 82.5 - 80.0 77.5 - 75.0 Within Among Within Among Within Among Within Among Yolla Bolly Trinity Alps Trinity Mtns. Marble Mtns. Fic. 3. Comparison of average within and among sub-regional Jaccard similarity coefficients for 15 foxtail pine stands in the Klamath Mountains. Differences are significant (ANOVA: F; 5, = 17.50, P = 0.0000009). N NM Northt or < North2 North3 North6 North? North15 Oe O Axis 1 V/ North3 North11 North13 North12 Ey o North10 Fic. 4. A Bray-Curtis ordination using Sorenson similarity coefficients showing patterns for 15 foxtail pine stands from the Klamath Mountains. Symbols are proportional to foxtail pine importance. Marble Mountains (closed dia- monds), Russian Peak (open triangles), Trinity Alps (open circles), Trinity Mountains (open squares), and Yolla Bolly Mountains (closed triangles). 2002] ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 41 TABLE 4. CHARACTERISTICS OF 15 FOXTAIL PINE STANDS IN THE SOUTHERN SIERRA NEVADA. Density given in trees/ hectare, basal area in m?/hectare, and relative values in percent. Data are from Vankat (1970) and Ryerson (1983). Averages +1 SE in parentheses. Vankat Stand Density Basal area 1 500 220 2 0 0 3 500 161 4 300 182 5) 600 67 6 400 66 1 200 97 8 600 n/a 2 200 151 10 — —: 11 — — 1192 — — 13 — — 14 — — 15 — — Average 367 (69) 118 (24) to 56 m*/ha (Table 4). Stand elevations ranged from 2900 m to 3650 m. Importance values for foxtail pine were at or approached 300 with western white pine, whitebark pine, and lodgepole pine as minor stand components. This prompted Vankat to iden- tify a foxtail pine forest type. Tree ring counts in- dicated that foxtail pine cover and density began increasing approximately 110 years ago in accor- dance with decreased grazing levels by sheep. Interregional comparison. A comparison to Ryerson’s data set showed that stand basal area, foxtail pine basal area, elevation, and conifer rich- ness differed between regions, but importance val- ues did not (Table 5). A comparison to Vankat’s data set showed that stand density, foxtail pine den- sity, and foxtail pine relative basal area differed be- tween regions (Table 5). Variances around these es- timates also differed between regions, with the southern Sierra Nevada having significantly greater variance estimates (Table 5). Comparisons between Ryerson’s and Vankat’s data sets showed differenc- es in stand density, foxtail pine relative density, and foxtail pine relative basal area (Table 5). The Bray-Curtis ordination using all stands did not cluster stands exclusively by region (Fig. 5), but there was broad separation of the majority of Klam- ath Mountain stands from a bifurcated grouping of southern Sierra Nevada stands dependent upon dominance of either foxtail or whitebark pine (Fig. 5). The stands with the highest importance values for foxtail pine were exclusively identified in the Sierra. Axis I in the ordination using all stands was highly correlated with foxtail pine importance (R = 0.855) and axis II with whitebark pine importance (R = —0.833). These axes had relatively high ex- planatory power (R? = 0.757). Four major clusters were identified: 1) stands dominated by foxtail Ryerson Foxtail pine Density Basal area importance 78 38 276 193 9 296 34 13 122 178 36 120 54 6 68 a2 18 90 74 N7/ 59 74 35 VS DAS 61 DW DDD 75 285 11 2 16 61 26 273 111 4 44 29 4 25) 51 15 181 100 (20) 24 (6) 145 (26) pine, 2) stands with foxtail pine and whitebark pine, 3) stands with foxtail pine and red fir, and 4) mixed stands with foxtail pine, red fir, and western white pine (Fig. 5). Stands with red fir and western white pine were associated with the lower elevations, while stands with whitebark pine were associated with higher elevations or northern stands. Incense-cedar, Doug- las-fir, and mountain hemlock were sampled only within the Klamath Mountains, while limber pine and mountain juniper (Juniperus occidentalis Hook.) were exclusively present in the Sierra Ne- vada. Red fir was present in the Sierra as the typical variety (Abies magnifica var. magnifica) and in the Klamath by Shasta red fir (Abies magnifica Andr. Murray var. shastensis Lemmon). An inverse rela- tionship between foxtail pine importance and co- nifer species richness was evident in the results of both procedures (Figs. 4, 6). DISCUSSION Recent genetic research has documented inter- regional and intraregional divergences of foxtail pine populations (Oline et al. 2000). We expected similar conclusions when compositionally compar- ing stands at these levels. Therefore we expected foxtail pine stands in close proximity within the Klamath Mountains to be similar, especially if they were located on the same substrate. Likewise, we expected the Klamath Mountains and southern Si- erra Nevada stands to differ in several ways. Both regions had simple stands dominated by foxtail pine, but this organization was found more often in the southern Sierra Nevada. Mixed stands with red fir or whitebark pine cut across regions. As found here, past studies suggested the lack of ecological differences between stands with different 42 MADRONO [Vol. 49 TABLE 5. STAND AND FOXTAIL PINE CHARACTERISTICS COMPARED USING TWO-SAMPLE T-TESTS (T), MANN-WHITNEY U TESTS (Z), AND HARTLEY'S EQUAL VARIANCE TESTS BETWEEN KLAMATH AND SIERRA REGIONS. Comparison to Ryerson (1983) is indicated by R and to Vankat (1970) by V. Equal variance tests were not conducted on non-normal data. Two-sample T and Mann-Whitney U tests Hartley’s test Variable Source Sample sizes Critical value P-value F-value P-value Conifer richness R n, = 15 T = 3.543** 0.00141** 1.600 0.389848 Elevation (m) R a s fe Z = 4.668** 0.000003** — — Importance value R a ic ET = —0:814 0.424904 4.026** 0.013294 Basal area R -, a a T = 1.847*** 0.08583*** 248.941 ** 0.000001 Density R = a ie T = 0.248 0.806775 264.552** 0.000001 Relative basal area R ze ie Z = —0.560 OD75511 — — Relative basal area* Vv : is Thy = PIS NEKO 0.003388** — — Relative density R ey z ie Z = —0.477 0.63325 a — Ny = 15 Relative density* Vv n, = 15 Tie eo nly lee 0.001521** — — Relative frequency R a a ie 1S) 02135 0.27096 Daten | 0.002213 Stand basal area R Fs is T= 22i0F= 0.036552** 264.552** 0.000001 Stand density R rf, B ie T = 0.848 0.405853 339582 0.029873 Stand density* Vv - = 13 Ti=s— 3.65338 0.004152** 4.039** 0.037774 ny = * Tests for statistical differences between these values from Ryerson (1983) and Vankat (1970) were significant with a = 0.05. ** Significant at a = 0.05. *** Significant at a = 0.10. red fir varieties (Barbour and Woodward 1985). Mixed foxtail pine stands with western white pine grew on sites with similar conditions to those for unmixed stands in both regions suggesting similar habitat needs by both species within and between regions. This result was expected because of west- ern white pine’s prevalence in many subalpine for- est types within the Californian mountains (Sawyer and Keeler-Wolf 1995). As expected there were species exclusive to each region (Griffin and Critchfield 1972), but they were not important in differentiating stands at the regional level. Ryerson found that mixed stands with whitebark pine were associated with less-developed soils at higher elevations typical of xeric, nutrient-limited sites, while mixed stands containing red fir were associated with deeper and more developed soils typical of mesic, nutrient-rich sites. Our analyses identified a cluster of foxtail pine-red fir dominated stands that cut across both regions. Within the Klamath Mountains, red fir was a more common associate of species rich stands, but these were nei- ther more mesic nor nutrient-rich than stands dom- inated by foxtail pine. Stands dominated almost ex- clusively by foxtail pine in the Klamath Mountains were typically found on sites intermediate to the extremes mentioned by Ryerson, on ultramafic soils, or homogeneous substrate compositions. An alternate explanation to Ryerson’s site limi- tation hypothesis may better explain the foxtail pine stands mixed with red fir. These stands may be the product of species-specific elevation limits. Within the Sierra Nevada, conifer species zone over a con- siderable elevation range. Much of the upper mon- tane red fir forests are replaced with lodgepole pine and whitebark pine forests as elevation increases (Potter 1998). Individual red fir trees are found in these types illustrating broad ecotones between for- est types. This is not the case in the compressed forest zones found in the Klamath Mountains. The comparison of Ryerson’s and Vankat’s data sets illustrated important differences suggesting that different sampling methods accounted for the dif- fering estimates of basal area. Stand selection cri- teria may have been even more important. Ryerson picked stands at marginal locations, while Vankat chose stands in areas with extensive foxtail pine dominance. The marginal stands of Ryerson were expected to have high variances if foxtail pine stands exhibited a core-periphery spatial structure. Comparisons reveled that Ryerson’s estimates were more variable and could be attributed to sampling from different core and peripheral populations. 2002] ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 43 Fic. 5. A Bray-Curtis ordination using Sorenson similarity coefficients showing patterns for 30 foxtail pine stands in the Klamath Mountains and southern Sierra Nevada. Open circles represent the Klamath Mountains stands. Closed triangles represent the southern Sierra Nevada stands. 1) stands dominated by foxtail pine, 2) stands with foxtail pine and whitebark pine, 3) stands with foxtail pine and red fir, and 4) mixed stands with foxtail pine, red fir, and western white pine. ABMA = red fir density, BA = stand basal area, PIAL = whitebark pine density, PIBA = foxtail pine density. Fic. 6. A Bray-Curtis ordination using Sorenson similarity coefficients showing patterns for 30 foxtail pine stands in the Klamath Mountains and southern Sierra Nevada. Open circles represent the Klamath Mountains stands. Closed triangles represent the southern Sierra Nevada stands. Symbol sizes are proportional to the conifer species richness found within each stand. 44 MADRONO The finding that called most for an interpretation was the inverse relationship between foxtail pine importance and conifer diversity prominent in the Klamath Mountains. Because conifer diversity was also correlated to substrate heterogeneity, simple causal explanations are difficult. Several authors have an explanation for this pattern that might be called the marginal hypothesis (Wright and Mooney 1965; Ryerson 1983; Sawyer and Thornburgh 1988; Kruckeberg 1992; Oline et al. 2000). They offer that within marginal habitats decreased biotic interactions lead to increased species diversity. Marginal habitats have been hypothesized as com- posed of nutrient limited soils (Sawyer and Thorn- burgh 1988; Kruckeberg 1992; Oline et al. 2000), extreme climate (Ryerson 1983), and cyclically dis- turbed habitat (Murray et al. 2000). Two possible marginal site types exist within the distributional limits of foxtail pine—high elevation sites and ultra- mafic soils. Ryerson proposed that the highest elevation stands within the Sierra Nevada simulated marginal habitat, but foxtail pine acts as a timberline species in the center of its range when south of whitebark pine’s range (Scuderi 1987). These stands are sim- ple with one or two species, rather than mixed as proposed by the marginal hypothesis. This may be a function of species distribution ranges along the Sierran crest. Similar elevations are absent in the Klamath Mountains, and elevation was not corre- lated to species richness. Ultramafic soils have been argued as marginal sites that isolate species and promote increased di- versity. This soil type is absent within the range of foxtail pine within the southern Sierra Nevada. Stands located on marginal ultramafic soils in this study were not more diverse than stands located at better sites (see Fig. 2). Nor were they less diverse with respect to conifer diversity. Stands with high conifer diversity were found on schists, gabbro, and peridotite in the Yolla Bolly Mountains, Trinity Alps, and Trinity Mountains, as were the stands dominated by foxtail pine. The most diverse stand, sampled at Russian Peak, was on glaciated granite. These results suggest that ultramafic soils do not directly control conifer species diversity as pro- posed in the marginal hypothesis. Nor do they con- trol foxtail pine importance (see Fig. 2). Conifer diversity in the Klamath Mountains was variable, and was greatest in the northern-most stands. The stands in the Yolla Bolly Mountains were well south of the range of lodgepole pine and whitebark pine and at the range limits for mountain hemlock and western white pine, reducing the spe- cies pool in the south. In the northern stands, these subalpine species mixed with montane conifers such as Douglas-fir, incense-cedar, and white fir. Within the Sierra Nevada, the limits of these mon- tane species are well below that of foxtail pine. Analysis of Jaccard similarity coefficients by sub-region reveled that foxtail pine stands in close [Vol. 49 proximity were more similar to one another than to stands in distant sub-regions. This pattern indicated an island-like distribution of foxtail pine and other species as predicted by the mountain island effect hypothesis (Brown 1971; Hamrick et al. 1994). This hypothesis argues that mountain tops are iso- lated collections of species experiencing ecological processes commonly observed within insular sys- tems. These patterns may be the result of slow dis- persal rates inherent to conifers and/or historical climate fluctuations. Mohr’s et al. (2000) recreation of post-glacial vegetation history at two lakes in the Trinity Mountains supports this possibility. The vegetation patterns and histories differed as much as they were similar even though these lakes are approximately 4.5 km apart. Inferences with our data suggest that the moun- tain island effect enhanced by substrate heteroge- neity better explains conifer species diversity pat- terns than does the marginal hypothesis. Substrate heterogeneity was positively correlated with conifer diversity and glaciated substrates. Rocky conditions such as moraines would isolate individual trees in favorable microsites if the species was in the area to take advantage of these sites. The high diversity stand at Crater Lake was partially logged and it had high diversity. Qualitative observations suggested that boulders isolated the new seedlings and sap- lings establishing after the disturbance. SUMMARY Foxtail pine stands within the Klamath Moun- tains and southern Sierra Nevada did not differ dra- matically in structure. Foxtail pine importance was constant across both regions, but species richness, stand basal area, and elevation differed between re- gions. Within the Klamath Mountains, conifer di- versity, foxtail pine relative density, and substrate heterogeneity differed among five sub-regions. Co- nifer diversity and foxtail pine importance were not correlated to substrate type (e.g., ultramafic soils) as previously hypothesized. Significant correlations were observed among foxtail pine importance, co- nifer diversity, and substrate heterogeneity. These correlations were more significant on glaciated sub- strates versus unglaciated substrates. Inferences from our data suggest that foxtail pine importance and conifer diversity in the Klamath Mountains may be regulated by a mountain island effect en- hanced through substrate heterogeneity, but more research is needed to tease apart the causal mech- anisms within these correlations. ACKNOWLEDGMENTS We would like to thank the Department of Biological Sciences at Humboldt State University, the Hayfork Rang- er District of Shasta-Trinity National Forest, and Linda Peak for use of sampling equipment. My fiancé Melissa L. Postler and coworker Vin D’ Angelo provided invalu- able field assistance, and Bob’s Auto and Tire Repair Ser- vice located in Red Bluff, California fixed numerous flat a 2002] tires obtained on the back roads of Siskiyou, Trinity, and Tehema counties. LITERATURE CITED BaiLey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Annals of the Mis- souri Botanical Gardens 57:210—249. BALL, J. T. 1976. Ecological survey of Last Chance Mead- ow Candidate Research Natural Area. Unpublished report. USDA Forest Service, Pacific Southwest Re- search Station, Albany, CA. BarsBour, M. G. AND R. A. WOODWARD. 1985. The Shasta fir forest of California. Canadian Journal of Forestry 15:570—576. BONHAM, C. D. 1989. Measurements for terrestrial vege- tation. John Wiley & Sons, New York. Brown, J. H. 1971. Mammals on mountaintops: Non- equilibrium insular biogeography. American Natural- ist 105:467—478. CRITCHFIELD, W. B. 1977. Hybridization of foxtail and bristlecone pines. Madrofio 24:193—211. GRIFFIN, J. R. AND W. B. CRITCHFIELD. 1972. The distri- bution of forest trees in California. Research Paper PSW-82. USDA Forest Service, Pacific Southwest Research Station, Albany, CA. Hamrick, J. L., A. E SCHNABEL, AND P. V. WELLS. 1994. Distribution of genetic diversity within and among populations of Great Basin conifers. Pp. 147—161 in K. T. Harper, L. L. St. Clair, K. H. Thorne, and W. M. Hess (eds.), Natural history of the Colorado Pla- teau and Great Basin. University of Colorado Press, Niwot, CO. HUTCHESON, K. 1970. A test for comparing diversities based on the Shannon formula. Journal of Theoretical Biology 29:151—154. KENT, M. AND P. COKER. 1992. Vegetation Description and Analysis: A Practical Approach. Belhaven Press, London. KRUCKEBERG, A. R. 1992. Plant life of western North American ultramafics. Pp. 31—74 in B. A. Roberts and J. Proctor (eds.), The ecology of areas with serpen- tinized rocks: A world view. Kluwer Academic Pub- lishers, Dordrecht, The Netherlands. KRusE, S. M. 1989. Climatic water budgets in the southern Sierra Nevada, California. Thesis, California State University, Fresno. Major, J. 1988. California climate in relation to vegeta- tion. Pp. 11—74 in M. G. Barbour and J. Major (eds.), Terrestrial vegetation of California (Expanded ver- sion), California Native Plant Society, Sacramento. MASTROGIUSEPPE, R. J. 1972. Geographic variation in fox- tail pine, Pinus balfouriana Grev. & Balf. Thesis, Humboldt State University. Arcata, CA. AND J. D. MASTROGIUSEPPE. 1980. A study of Pi- ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 45 nus balfouriana Grev.& Balf. (Pinaceae). Systematic Botany 5:86—104. McCune, B. 1993. PC —Ord. Version 3.11 computer pack- age. Mone, J. A., C. WHITLOCK, AND C. N. SKINNER. 2000. Postglacial vegetation and fire history, eastern Klam- ath Mountains, California, USA. The Holocene 10: 587-601. Mirovy, N. T. 1967. The Genus Pinus. The Ronald Press Company, New York. Murray, M. P.,, S. C. BUNTING, AND P. MorGAn. 2000. Landscape trends (1753—1993) of whitebark pine (Pi- nus albicaulis) forests in the West Big Hole Range of Idaho/Montana, U.S.A. Arctic, Antarctic, and Alpine Research 32:412—418. OLINE, D. K., J. B. MITTON, AND M. C. GRANT. 2000. Pop- ulation and subspecific genetic differentiation in the foxtail pine (Pinus balfouriana). Evolution 54:1813-— 1819. POLLARD, J. H. 1971. On distance estimators of density in randomly distributed forests. Biometrics 27:991— 1002. Potter, D. A. 1998. Forested communities of the upper montane in the central and southern Sierra Nevada. General Technical Report PSW-GTR-169, USDA Forest Service, Pacific Southwest Research Station, Albany, CA. RAVEN, P. H. AND D. I. AXELROD. 1978. Origin and rela- tionships of the California flora. University of Cali- fornia Publications in Botany 72:1—134. RYERSON, D. 1983. Population structure of Pinus balfour- iana Grev. & Balf. along the margins of its distribu- tion area in the Sierran and Klamath Regions of Cal- ifornia. Thesis, Sacramento State University. Sacra- mento, CA. SAWYER, J. O. AND D. THORNBURGH. 1988. Montane and subalpine vegetation of the Klamath Mountains. Pp. 699-732 in M. G. Barbour and J. Major (eds.), Ter- restrial vegetation of California, Expanded ed., Cali- fornia Native Plant Society, Sacramento. AND T. KEELER-WOoLF. 1995. A Manual of Cali- fornia Vegetation. California Native Plant Society, Sacramento. SCUDERI, L. 1987. Late-Holocene upper timberline varia- tion in the southern Sierra Nevada. Nature 325:242— 244. VANKAT, J. L. 1970. Vegetation change in Sequoia Na- tional Park, California. Dissertation, University of California, Davis. WALKER, R. B. 1954. Factors affecting plant growth on serpentine soils. Ecology 35:259—266. WRIGHT, R. D. AND H. A. Mooney. 1965. Substrate-ori- ented distribution of bristlecone pine in the White Mountains of California. The American Midland Nat- uralist 73:257—284. ZAR, J. H. 1999. Biostatistical Analysis. Prentice Hall, Up- per Saddle River, NJ. MApRONO, Vol. 49, No. 1, pp. 46-47, 2002 REVIEW The manzanitas of California, also of Mexico and the world, by Phillip V. Wells. 2000. Published by P. V. Wells, Department of Ecology and Evolution- ary Biology, Haworth Hall, University of Kansas, Lawrence KS 66045. Available from Cody’s Books, Berkeley, CA for $53.00. ISBN: 0-933994- DD. The recent review of Trees and Shrubs of Cali- fornia (Stuart and Sawyer 2001) by Rejmanek (2001) underscores the need for a review of The manzanitas of California (Wells 2000). Rejmanek points out that there are “‘more than 40 excluded species of Arctostaphylos” in Stuart and Sawyer (2001) and he suggests that people would be better off getting Wells (2000) if they have an interest in this quintessentially Californian group of woody shrubs. To a degree, we agree—but with some im- portant caveats that should be kept in mind. P. V. Wells has made the study of manzanitas a lifetime work and this self-published, un-peer re- viewed book demonstrates both the best and the worst of this kind of situation. Most of the infor- mation in the book, for example, can be found in previous publications of his, including most of the figures (See, e.g., Ecological Monographs 32:79-— 103 [1962], Evolution 23:264—267 [1969], The Four Seasons 7:17—21 [1987], 8:46—70 [1990], 9: 64-69 [1992]). Wells published his treatment on Arctostaphylos in the 1993 Jepson Manual (Hick- man 1993) (Chapter V). He also published a phy- logenetic hypothesis for Arctostaphylos in 1992 (Chapter IV) (Wells 1992) in which he divides the genus into two subgenera and six sections. This lat- est book presents this treatment in detail and ig- nores later publications that cast serious doubt on Wells’ phylogenetic hypothesis (Markos et al. 1998). He then goes on to trivialize important new findings in the genus by Keeley and his students (Keeley 1994; Keeley et al. 1997a, b). Figures in this book come from the original publications and many of the species names are woefully out of date, no longer valid even in the treatment by Wells. In Chapter V, he includes small, fuzzy, black and white photos of herbarium specimens to illustrate species; none of these are sharp enough or show the appropriate characters to be of much use. While he provides some interesting discussions, for ex- ample, concerning reticulate evolution, Wells has not published data that would enable objective evaluation of his conclusions, nor does it exist as a table or appendix in this book. The overall impres- sion is that Wells has decreed an ideal manzanita world that he can perceive and which now has been formally revealed. Indeed, but if only those plants in the field would behave! With these minor criticisms aside, we acknowl- edge that the majority of the taxa recognized by Wells are probably distinct lineages and this vol- ume provides a wealth of information about them. Indeed, we agree with Wells that manzanitas are the most diverse and fascinating genus of woody shrubs in the California Floristic Province (one of 25 global “‘hotspots’’ on the planet [Mitermeier et al. 2000]). Most importantly, this book distills Wells’ long years of scholarly research into the ge- nus and provides an invaluable resource for any serious manzanita student. As mentioned above, Wells develops an intriguing chapter on reticulate evolution in Arctostaphylos (Chapter VII). This in- formative discussion concerning hybridization and speciation in manzanitas articulates his hypothesis concerning diploid hybridization. Wells presents scatter diagrams and multivariate-analytic figures that suggest evidence of hybrid origin between spe- cies such as A. canescens and A. andersonii (lead- ing to intermediate species such as A. glutinosa and A. auriculata). Unfortunately, the data supporting these studies is missing, and in tables speculating on species of possible hybrid origin, the inclusion of many plausible parents (e.g., A. glutinosa from A. canescens X A. andersonii) are undermined by a number of frankly preposterous suggestions (e.g., A. pilosula from A. wellsii X A. glauca). Wells also provides a lot of practical material. He gives interesting regional keys to manzanitas in Chapter VI, a chapter rich in historical lore. The Introduction contains a discussion of generic rela- tionships between Arctostaphylos and other mem- bers of the Arbutoideae (Arbuteae) distilled from Diggs and Breckon (1981) through Wells’ own per- spective. Hileman et al. (2001) provide a molecular counterpoint that reinforces the general accuracy of the generic circumscriptions that Wells describes for this monophyletic group. The Character Analysis (Chapter II) of Arcto- staphylos illustrates the fundamental flaw in Wells’ classification system. He lists 70 morphological traits that he claims he has analyzed for 61 species. He then describes character states for these traits and goes on to fashion elaborate descriptions of species that are infinitely complex based upon this enormous data set (again, which is not tabulated in the book). These species descriptions are enumer- ated in Chapter IV and ordered into the same phy- logenetic scheme that Wells (1992) proposed eight years ago. The flaw is that manzanitas don’t obey the world according to P V. Wells, as was illus- trated by Keeley (1994), Markos et al. (1998), Va- sey and Parker (1999), and our extensive field ob- servations. Wells continues with the belief that pan- icles can be described as racemes (with up to 7 2002] branches!), and that nascent inflorescences with variable bract characters (leafy vs. scaly bracts) can be classified as either one state or the other exclu- sively. As a consequence, we have found that a great deal of confusion and mystification is created by a treatment that tends to elicit more frustration than enlightenment. And so, we caution, if you buy this book, focus on the real entities and not necessarily on the bi- or trinomial classification system offered by Wells. Luxuriate in the lore but take lightly the conclu- sions. Particularly, defer judgments regarding phy- logenetic relationships until an alternative and probably molecular phylogeny is worked out that meshes with morphology, cytology, and ecological information and that will also employ a more con- temporary species concept. There is a rich literature on manzanitas by some great California botanists and evolutionary biologists (e.g., Eastwood [1934], Jepson [1922], Dobzhansky [1953], Stebbins and Major [1965], etc.). These scientists recognized the extraordinary importance of Arctostaphylos to un- derstanding driving forces in the evolution of Cal- ifornia’s remarkable flora. Wells’ book culminates a wild and woolly era in the history of manzanita taxonomy that diverged from this venerable tradi- tion. Will Wells (2000) be the last word on man- zanitas? We don’t think so (Keeley 1998; Keeley et al. 1994, 1997a, b; Markos et al. 1998; Vasey and Parker 1999; Hileman et al. 2001). —MICHAEL C. VASEY AND V. THOMAS PARKER, Depart- ment of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, CA 94132. mvasey @ sfsu.edu; parker@sfsu.edu LITERATURE CITED Diccs, G. M. AND G. J. BRECKON. 1981. Generic circum- scription in the Arbuteae (Ericaceae). In G. M. Diggs, Systematic studies in the Arbuteae. Ph.D. dissertation. University of Wisconsin, Madison, WI. DOBZHANSKY, T. 1953. Natural hybrids of two species of REVIEW 47 Arctostaphylos in the Yosemite region of California. Heredity 7:73—79. Eastwoop, A. 1934. A revision of Arctostaphylos with key and descriptions. Leaflets of Western Botany 1: 105-127. HICKMAN, J. C. (ed.). 1993. The Jepson manual: Higher plants of California. University of California Press, Berkeley. HILEMAN, L. C., M. C. VASEy, AND V. T. PARKER. 2001. Phylogeny and biogeography of the Arbutoideae (Er- icaceae): Implications for the Madrean-Tethyan hy- pothesis. Systematic Botany 26:131—143. JEPSON, W. L. 1922. Revision of Californian Arctostaphyli. Madrono 1:78—86. KEELEY, J. E. 1994. Arctostaphylos rainbowensis, a new burl-forming manzanita from northern San Diego County, California. Madrono 41:1—12. , L. BOYKIN, AND A. MASssIHI. 1997a. Phenetic analysis of Arctostaphylos parryana: (1) Two new burl-forming subspecies. Madrono 44:253—267. , A. MAssiHt, J.D. RODRIGUEZ, AND S.A. HIRALES. 1997b. Arctostaphylos incognita, a new species and its phenetic relationship to other manzanitas of Baja California. Madrofio 44:137—150. MARKOs, S., L.C. HILEMAN, M.C. VASEY AND V.T. PARKER. 1998. Phylogeny of the Arctostaphylos hookeri com- plex (Ericaceae) based on nrDNA data. Madrono 45: 187-199. MITERMEIER, R.A., N. MEYERS, C.G. MITERMEIER, AND N. MEYERS. 2000. Hotspots: Earth’s biologically richest and most endangered terrestrial ecosystems. Univer- sity of Chicago Press, Chicago, IL. REJMANEK, M. 2001. Trees and shrubs of California by John D. Stuart and John O. Sawyer: A book review. Madrono 48:128—129. STEBBINS, G. L. AND J. Mayor. 1965. Endemism and spe- ciation in the California flora. Ecological Monographs 35:79-102. STUART, J. D. AND J. O. SAwYER. 2001. Trees and shrubs of California. University of California Press, Berke- ley. VASEY, M. C. AND V. T. PARKER. 1999. Nascent inflores- cences in Arctostaphylos pringlei: response to Keeley and Wells. Madrofo 46:51—54. WELLS, P. V. 1992. Subgenera and sections of Arctostaph- ylos. The Four Seasons 9:64—69. . 2000. The manzanitas of California, also of Mex- ico and the world. Published by the Author. MADRONO, Vol. 49, No. 1, p. 48, 2002 Illustrated field guide to selected rare plants of northern California. Edited by Gary Nakamua and Julie Kiersteand Nelson. 2001. University of Cali- fornia Agriculture and Natural Resources Publica- tion 3395, Oakland, CA. 370 pp. Softcover $36.00. ISBN 1-879906-5470. For those interested in the botany of northern California, this field guide is a real treat. This book richly illustrates and describes 149 of the rarer plant taxa found in northern 10 counties in the state (Butte, Del Norte, Glenn, Humboldt, Lassen, Mo- doc, Plumas, Shasta, Siskiyou, and Trinity Coun- ties). The authors focus mainly on plants presumed extinct or rare in California, Lists 1A and 1B re- spectively in the 6th edition CNPS Inventory (CNPS 2001). Also included are six List 2 (rare in California, but more common elsewhere), one on List 3 (review list), and four on List 4 (watch list). The 149 taxa include three that are state-listed as endangered, 12 that are state-listed as rare, eight that are federally-list as endangered, and three that are federally listed as threatened. It is easy to jump over the first 40 pages of in- troduction to enjoy the species descriptions. Each is a two page treatment. The spiral binding allows for quick access. Plants are arranged alphabetically by genus, with the name in the upper left and lower right corners to make it easy to find a plant. Each treatment involves a large photograph of a mature plant, a small photograph of its habitat, and a map indicating occurrences by USGS quadrangle loca- tion. On the facing page, a line art illustration accom- panies a description of the plant, habitat, and lo- cation. Scientific names follow the new edition of the CNPS Inventory, as do synonymy, common name(s), family names, distribution, elevation, and quadrangle codes. Habitat designations generally follow those in the CNPS Inventory, but in some cases, the habitat descriptions are broader. In the Key Feature section, the first paragraph typically describes the plant. The second paragraph discusses similar taxa, and, if necessary, instructs the reader to “‘consult an expert to verify identification.” If this is the case, a list of diagnostic features accom- panies an expert symbol. Flowering times and iden- tification times, which may differ if fruits are re- quired, finish the presentation. Illustrations are drawn from 26 sources including lovely, original art by Linda Vorobik. The book has no keys, exhaustive descriptions, or complete synonymy. It assumes a basic knowl- edge of plant identification and is not intended to substitute for standard botanical references or field guides. Instead, the book’s purposes are to “. help the reader develop an accurate search image ” and to “... learn how to accurately distin- guish rare plants from similar species in the field .. The editors involved 28 contributors, who make up the Northern California Botanists, an ad hoc committee of federal, state, and consulting bot- anists. More than other botanists, they are faced each field season with a new set of recruits to con- duct plant surveys. This book is designed for them, but it will be useful to seasoned botanists as well. Turning to the introduction, the reader finds the expected definitions, a short description of state and federal laws concerning protecting and conserving plants, and an explanation of how to use guide. Next come two extensive tables. The first lists spe- cies by geographic subdivisions of the state found in The Jepson Manual (Hickman 1993). The second table list species by habitat in each region. These tables are enlightening. As expected, rare plants along the coast were most common on the dunes, but the north coast conifer forest is not far behind. I expected to find a spate of serpentine spe- cies in the Klamath and North Coast Ranges, but not necessarily in mid-elevation forests and wood- lands. The same conclusion came from reviewing the lists for the Cascades, Sierra Nevada, and the Great Basin. The point was made most vividly when I saw the habitats of Cryptantha crinita and Lotus rubriflorus. The traveler on Interstate 5 drives by miles of similar looking dry streams and grass- lands. This book will help people get over the idea that all rare plants only grow in special places. With this book in hand and its great photographs, people can shed that misconception, as well as develop accurate search images for northern California’s rare plants. I doubt that you will find the book in most book- stores, but it can be ordered from the University of California, Agriculture and Natural Resources (ANR) Catalog at http://anrcatalog.ucdavis.edu/ 15 —JOHN O. SAWYER, Department of Biological Sciences, Humboldt State University, Arcata, CA 95521. LITERATURE CITED CNPS. 2001. Inventory of rare and endangered plants of California, 6th ed. Rare Plant Scientific Advisory Committee, David P. Tibor, Convening Editor. Cali- fornia Native Plant Society. Sacramento HICKMAN, J. C. (ed.). 1993. The Jepson manual: Higher plants of California. University of California Press, Berkeley. MApDRONO, Vol. 49, No. 1, pp. 49-53, 2002 NOTEWORTHY BRYOPHYTE RECORDS FROM THE MOJAVE DESERT LLoyD R. STARK Department of Biological Sciences, University of Nevada, 4505 Maryland Parkway, Las Vegas, NV 89154-4004 ALAN T. WHITTEMORE! Missouri Botanical Garden, PO. Box 299, St. Louis, MO 63166 BRENT D. MISHLER University Herbarium, Jepson Herbarium, and Department of Integrative Biology, 1001 Valley Life Sciences Bld., #2465, University of California, Berkeley, CA 94720-2465 ABSTRACT Significant range expansions in the northern Mojave Desert are documented for twenty-three species of bryophytes, nine of which are new to the region. Barbula convoluta, Claopodium whippleanum, En- tosthodon planoconvexus, Fissidens sublimbatus, Grimmia americana, Reboulia hemispherica, Syntrichia bartramii, Weissia condensa, and Asterella californica are new to the flora of Nevada. The northern Mojave Desert encompasses four counties: Clark and Nye Cos., Nevada, Washington Co., Utah, and Mohave Co., Arizona (MacMahon and Wagner 1985). Recently, a list of bryophyte species from this region was compiled from the lit- erature (Stark and Whittemore 2000), drawing chief- ly from state and regional checklists and treatments (Lawton 1958; Haring 1961; Flowers 1973; Spence 1988). The bryophyte flora of the region consists of 75 species of mosses and 5 species of liverworts. More than half of the bryophytes from the northern Mojave Desert belong to the more xeric families Pottiaceae, Grimmiaceae, and Orthotrichaceae. Ongoing collecting efforts in southern Nevada indicate that this region is not well explored bryo- logically, and this applies to the entire state (Heise 2000). The recent discovery of a new species of Didymodon in the remote southeastern portion of Nevada (Zander et al. 1995), and an as yet unde- scribed species in the liverwort genus Targionia that appears to be endemic to the Mojave Desert (Whittemore 1996) indicate exploration is needed. In the present paper, we discuss several species of bryophytes that are reported new to the northern Mojave Desert, or whose distributions are consid- erably broadened within the region. Nomenclature follows Anderson et al. (1990), Zander (1993), and Stotler and Crandall-Stotler (1977). BRYOPHYTA Barbula convoluta Hedw. Nevada, Clark Co., southern Gale Hills, lower end of Lovell Wash, 0.3 km upstream of confluence ' Present address: U.S. National Arboretum, 3501 New York Avenue NE, Washington, DC, USA 2002-1958 of Lovell Wash and West End Wash, along steep, north-facing outcrop adjacent to dirt road, elev. 550 m, Stark NV-1941 (UNLV), 1942 (UNLV, BUF). Nevada state record. This is the first report of this rather wide-ranging species from the Mojave Des- ert, with the only other report from the interior ba- sins of North America (southern Idaho, Flowers 1973; Spence 1988). From western North America, B. convoluta is known from British Columbia to Baja California (Lawton 1971; Zander 1994a). Claopodium whippleanum (Sull. in Whipple & Ives) Ren. & Card. Nevada, Clark Co., Spring Mountains, Red Rock Canyon National Recreation Area, steep side can- yon near mouth of Red Rock Canyon, near conflu- ence with Red Rock Wash; in deep shade beneath boulders, on sandstone rock and dead wood, elev. 1450 m, Stark NV-316 (UNLV). Nevada state rec- ord. Found in an area that never receives direct sun- light, in a side canyon on a steep slope under boul- ders. In North America, the species ranges from northwestern Mexico to British Columbia (Crum and Buck 1994), and is reported from southern Cal- ifornia as occasional in cismontane lowlands (Har- thill et al. 1979). It is distributed in western North America and also the Mediterranean region (Scho- field and Crum 1972). A disjunct population was reported from high elevation in northeastern Ari- zona (Apache Co., 9500 ft, Haring 1961). Coscinodon calyptratus (Hook. in Drumm.) C. Jens. ex Kindb. Reported from the Mojave Desert of southwest- ern Utah (Hastings 1999). Previously reported from Mohave County, Arizona (Haring 1961, as Grim- mia calyptrata Hook.), and from Lincoln Co., Ne- 50 MADRONO vada (Lawton 1958, as Grimmia calyptrata Hook.). This species is broadly distributed in Nevada and Utah, reaching its southwesternmost extent in the Mojave Desert (Hastings 1999). Crossidium seriatum Crum & Steere Nevada, Clark County, northern foothills of Lime Ridge, ca. 16 km north of Gold Butte, elev. 487 m, Stark NV-232a (UNLV, MEXU); southern Moapa Valley, sandstone bluffs along periphery of valley, ca. 8 km south of Overton, along Hwy 169, elev. 488 m, Stark NV-872 (UNLV, MEXU); Black Mountains, Lake Mead National Recreation Area, gypsum formation 2.1 km down Boathouse Cove Road from North Shore Road turnoff, elev. 650 m, Stark & Bonine NV-3045 (UNLV); Arizona, Mo- have County, Lake Mead National Recreation Area, Lake Mead Landing, mouth of Kingman Wash, elev. 396 m, W. Niles s.n., 24 Feb 1995 (UNLV). Recently recorded from the state of Nevada (Zander et al. 1995), the range of C. seriatum is expanded to a scattering of southern Nevada locations and also into Mohave County, Arizona. This globally rare species is presently known from only a handful of populations worldwide outside of the state of Nevada: Mariposa County, Arizona (Zander 1977), Cedros Island, Mexico, San Diego County, Cali- fornia (Stark and Whittemore 1992), and Spain (Cano et al. 1992). Didymodon vinealis (Brid.) Zand. Nevada, Clark County, Southern Gale Hills, low- er end of Lovell Wash, 0.3 km upstream of conflu- ence of Lovell Wash and West End Wash, along steep, north-facing outcrop adjacent to dirt road, elev. 550 m, Stark NV-1946 (UNLV); Lake Mead National Recreation Area, 4.3 km south of Rogers Spring, narrow canyon on north side of North Shore Rd, elev. 750 m, Stark NV-2059a (UNLV); Newberry Mountains, Lake Mead National Recre- ation Area, “‘Needles Eye’’, ca. 7.2 km north on Christmas Tree Pass Rd from Hwy 77E, elev. 817 m, Stark NV-76 (UNLV). Recently reported from the northern Mojave from the River Mountains as an incidental species (Stark 1997), the range of D. vinealis is considerably broadened here. Entosthodon planoconvexus (Bartr.) Grout Nevada, Nye County, Nevada Test Site, Rock Valley, north-facing foothills of Spectre Range, elev. 1159 m, growing with an undescribed species of Targionia, Stark NV-724 (UNLV). Nevada state record, and one of only four localities worldwide. Other known localities include Washington County, Utah (Flowers 1973), Pima County, Arizona (Har- ing 1961), and the northern Egyptian desert (Shab- bara 1999). The specimen cited herein (Stark NV- 724) differs from descriptions of E. planoconvexus in having a pale (not red) seta, a relatively narrow capsule that is strongly contracted under the mouth [Vol. 49 when dry, and a rudimentary peristome. The latter features align it with E. tucsonii (Bartr.) Grout. However, E. tucsonii has spores twice as large as Stark NV-724, making this determination improb- able. Clearly, a revision of North American Entos- thodon is needed. Fissidens sublimbatus Grout Nevada, Clark County, Newberry Mountains, Lake Mead National Recreation Area, Pipe Spring Canyon, near Pipe Spring, west-facing soil in rock crevice, elev. 732 m, Stark NV-99 (UNLV, PAC). New to the northern Mojave Desert and a Nevada state record. Known also from Arizona, California, New Mexico, and Baja California (Pursell 1994). Funaria hygrometrica Hedw. Nevada, Clark County, Newberry Mountains, Lake Mead National Recreation Area, ‘‘Needles Eye,” ca. 7.2 km north on Christmas Tree Pass Rd from Hwy 77E, elev. 817 m, Stark NV-SO (UNLV); Virgin Mountains, east base of South Virgin Peak Ridge, Lime Spring Canyon, abundant in moist drainages, W. Niles s.n., 22 Mar 1996 (UNLYV). This cosmopolitan species is reported as new to southern Nevada, having previously been reported from Mohave County, Arizona (Haring 1961). Funaria muhlenbergii Turn. Nevada, Clark County, northern River Moun- tains, Lake Mead National Recreation Area, 5.6 km due east of Saddle Island, elev. 610 m, Stark NV- 144 (UNLV); Eldorado Mountains, Keyhole Can- yon Archeological Site, just north of base of Key- hole Canyon, elev. 274 m, Stark NV-190b (UNLY); Muddy Mountains, southern end of White Basin, adjacent to West Longwell Ridge, ca. 6.4 km by road northwest of Bitter Spring, elev. 650 m, Stark & Bonine NV-3013 (UNLV). Reported from the California Mojave (Harthill et al. 1979), and broad- ly distributed in western North America (Smith 1994). Grimmia americana Bartr. Nevada, Clark County, Newberry Mountains, Lake Mead National Recreation Area, Grapevine Canyon, on north-facing rock, elev. 793 m, Stark NV-16 (UNLV). Nevada State record, and one of only three populations of this species known world- wide, one from Arizona (Pima Co., Crum 1994), and one from western Texas (type locality, Jeff Da- vis County, Texas; Crum and Anderson 1981). This species is not listed in Haring (1961) as occurring in Arizona, and is not yet known from Mexico (Crum 1994). Grimmia americana has bistratose upper leaf cells, which distinguish it from G. pla- giopodia Hedw., and is peristomate, which distin- guishes it from G. anodon Bruch & Schimp. in B.S.G. Grimmia plagiopodia is listed in Haring 2002] (1961) as occurring only in Yavapai County, Ari- zona, from two collections, while G. anodon 1s widespread in the southwestern U.S. (Lavin 1982). Grimmia moxleyi Williams in Holz. Nevada, Clark County, Muddy Mountains, southern end of White Basin, adjacent to West Longwell Ridge, ca. 6.4 km by road northwest of Bitter Spring, elev. 650 m, Stark & Bonine NV- 3004 (UNLV); foothills of Black Mountains, Lake Mead National Recreation Area, along tributary to Manganese Wash (from the northwest), ca. 6.4 km north of Boathouse Cove, elev. 650 m, Stark & Bonine NV-3071 (UNLV). New to the northern Mo- jave Desert exclusive of California. Grimmia mox- leyi is reported from the California Mojave Desert as infrequent (Harthill et al. 1979), and its presence is expected based on a Death Valley locality noted in Koch (1954). A common associate of G. orbi- cularis Bruch ex Wils. in Nevada, G. moxleyi is endemic to the southwestern U.S. and adjacent northern Mexico (Greven 1999). It has been re- ported without specifics from Arizona and Nevada (Crum 1994). Recently, Munoz (2000) regarded G. moxleyi aS synonymous with the wide-ranging G. orbicularis, indicating ample variability in hair- point development among southwestern popula- tions. Homalothecium nevadense (Lesq.) Ren. & Card. Nevada, Nye County, Spring Mountains, Wood Canyon, in vicinity of Wood Canyon Spring, Stark & Landau NV-1794 (UNLV). First report for the Mojave Desert. This species is reported from south- ern California (Harthill et al. 1979) as frequent in the region, but excluding deserts. The species reaches its southernmost extent in northern Arizona (Flowers 1973). Hypnum vaucheri Lesq. Nevada, Clark County, Spring Mountains, Red Rock Canyon National Recreation Area, steep side canyon near mouth of Red Rock Canyon, near con- fluence with Red Rock Wash, on shaded rock, elev. 1450 m, Stark NV-315b (UNLV). New to the Mo- jave Desert. Frequent in southern Utah along the Colorado-Green River Basin (Flowers 1973). Microbryum starkeanum (Hedw.) Zand. Nevada, Clark County, lower end of Borax Wash, southern Gale Hills, Stark NV-1944 (UNLV). New to the Mojave Desert. Guerra and Cano (2000) prefer to retain this species in Pottia, as Pottia starckeana (Hedw.) Miill., because of its stegocar- pous sporophyte. Pseudocrossidium crinitum (Schultz) Zand. Nevada, Clark County, Muddy Mountains, Val- ley of Fire State Park, 0.5 km south of ‘“‘Mouse’s STARK ET AL.: MOJAVE DESERT BRYOPHYTES 51 Tank,’’ sandstone bluffs on west side of road, ele- vation 700 m, Stark & Bonine NV-3076, 3084 (UNLV). Second report for state of Nevada. The locality cited may be the same populations noted by Lawton (1958, as Tortula aurea Bartr.). This report thus confirms the only known population from the Mojave Desert, with the nearest known locality just outside the Mojave Desert in southern Utah (Spence 1987). This species is fairly common in Mexico, according to Zander (1994b), occurring in several Mexican states, and is listed from five counties in Arizona (Haring 1961). Despite the abundance of P. crinitum at this site, no sporo- phytes were found, consistent with the pattern of an absence of male plants in North America. Pterygoneurum subsessile (Brid.) Jur. Nevada, Clark County, Eldorado Mountains, Keyhole Canyon Archeological Site, just north of base of Keyhole Canyon, elev. 274 m, Stark NV- 192a (UNLV); Newberry Mountains, Lake Mead National Recreation Area, Grapevine Canyon, 3.2 km north on Christmas Tree Pass Rd from Hwy 77E, beyond petroglyphs to the east, elev. 854 m, Stark NV-22c (UNLV). Second report from the northern Mojave Desert; found previously in Wash- ington County, Utah (Flowers 1973). Syntrichia bartramii (Steere in Grout) Zand. Nevada, Clark County, Newberry Mountains, Lake Mead National Recreation Area, Grapevine Canyon, 3.2 km north on Christmas Tree Pass Rd from Hwy 77E, beyond petroglyphs to the east, elev. 854 m, Stark NV-32B, 38B (UNLV). Nevada state record. Previously reported from the southern Californian Mojave Desert (Harthill et al. 1979), S. bartramii occurs in northwestern Mexico and in the bordering states of Arizona, New Mexico, and Tex- as, with infrequent reports north of this region (Mishler 1994). One of the two specimens cited above from Nevada was found on juniper growing with S. pagorum, noteworthy in that epiphytic mosses are exceedingly rare in the Mojave Desert below 2000 m. Variation in this species is complex and should be studied in association with plants oc- curring in southern California, where it is possible that an undescribed species is present. Syntrichia pagorum (Milde) Amann Nevada, Clark County, Newberry Mountains, Lake Mead National Recreation Area, “‘Needles Eye,” ca. 7.2 km north on Christmas Tree Pass Rd from Hwy 77E, elev. 817 m, Stark NV-82A (UNLV); Pipe Spring Canyon, in vicinity of Pipe Spring, elev. 732 m, Stark NV-91 (UNLV). New to the Mojave Desert. Previously reported from Ne- vada (Crum and Anderson 1981), this species is distributed from eastern North America across the southern portion of the USA to the west coast. However, reports are lacking for Utah and southern 52 MADRONO California. Syntrichia pagorum is known only from female plants in the USA. Syntrichia princeps (De Not.) Mitt. Nevada, Clark County, Spring Mountains, Red Rock Canyon National Recreation Area, near mouth of Red Rock Canyon, near confluence with Red Rock Wash, on partially shaded rock, elev. 1450 m, Stark NV-291 (UNLV); Virgin Mountains, east base of South Virgin Peak Ridge, Lime Spring Canyon, on north-facing slope, at edge of water- course on moist drainages, W. Niles s.n., 22 March 1996 (UNLV). These two localities constitute the second and third reports from the northern Mojave Desert, with the first from Washington County, Utah (Flowers 1973). Syntrichia princeps 1s distin- guished from the related S. ruralis (Hedw.) Web. & Mohr by its (often) synoicous condition, and is dis- junct from the western coast of North America to the Spring Mountains and Virgin Mountains of southern Nevada and southern Utah, respectively. Syntrichia princeps is one of several species in Utah known only from Washington County (Flow- ers 1973). Syntrichia ruralis (Hedw.) Web. & Mohr Nevada, Muddy Mountains, Valley of Fire State Park, 0.5 km south of ‘““Mouse’s Tank,’’ sandstone bluffs on west side of road, elevation 700 m, Stark & Bonine 3077, 3078 (UNLV). Oddly, S. ruralis is known from only two sites from the northern Mo- jave Desert, from Mohave County, Arizona (Haring 1961), and from the River Mountains in Nevada (Stark et al. 1998). The locality given represents one of the few sites known to the authors in the Mojave Desert where male, female, and sporo- phytic plants co-occur. Tortula atrovirens (Sm.) Lindb. Nevada, Clark County, Bitter Spring Valley, Echo Wash, gypsum formation ca. 1.6 km east of Bitter Spring, elev. 530 m, Stark NV-2087 (UNLV); northern River Mountains, Lake Mead National Recreation Area, 5.6 km due east of Saddle Island, elev. 610 m, Stark NV-139a (UNLV); Newberry Mountains, Lake Mead National Recreation Area, Grapevine Canyon, 3.2 km north on Christmas Tree Pass Rd from Hwy 77E, beyond petroglyphs to the east, elev. 854 m, Stark NV-39a (UNLV). Only a single prior report exists from the northern Mojave Desert exists, and this as an incidental (Stark et al. 1998), thus indicating that the species is probably much more common than reports indicate. Tortula atrovirens was previously known as Desmatodon convolutus (Brid.) Grout. Weissia condensa (Voit) Lindb. Nevada, Clark County, foothills on northwest side of River Mountains, ca. 6.4 km from down- [Vol. 49 town Henderson, elev. 671 m, Stark NV-120 (UNLV, BUF). First report from southern Nevada and probably a state record. Reported from Arizo- na, southern Utah, Texas, and Mexico, W. condensa is a widespread species that also occurs in South America, Africa, and Europe (Flowers 1973, as W. tortilis (Schwaegr.) C. Muell.; Zander 1994c). HEPATICOPHYTA Asterella californica (Hampe) Underw. Nevada, Clark County, Lake Mead National Rec- reation Area, Indian Hills, north of Devil’s Cove, Gold Butte area, T20S, R7OE, at base of limestone boulders, elevation 650 m, W. E. Niles s.n., 6 March 1998 (UNLV); Spring Mountains, Red Rock National Conservation Area, near mouth of Red Rock Canyon near confluence with Red Rock Wash, T20S, R58E, S32, damp shaded soil beneath overhang at base of north-facing cliff, elev. 1450 m., A. T. Whittemore 68587 (MO). Nevada state rec- ord. Known from Mohave Co., Arizona (Evans 1917) and from Riverside Co., CA, near Palm Springs (S. B. Parish 3890, CAS). Reboulia hemispherica (L.) Raddi Nevada, Clark County, Spring Mountains, Red Rock National Conservation Area, near Willow Springs, T21S, R58E, S5, shaded gorge, in cracks in cliff, elev. 1500 m, A. 7. Whittemore 6891 (MO). Nevada state record. This is the first report of this rather wide-ranging species from the Mojave Des- ert. Reboulia hemispherica is fairly common in New Mexico and eastern Arizona, but it is rare and local west of these states, currently known only from two collections from the northern Sierra Ne- vada in California and a few scattered sites in the Pacific northwest. ACKNOWLEDGMENTS We thank the National Geographic Society (grant no. 5429-95) for providing funds for travel and the Missouri Botanical Garden for administering this grant; the U.S. Bureau of Land Management and the National Park Ser- vice (Lake Mead) for funding support; Wes Niles for con- tributing specimens critical to this study; Mary Bonine, Fred Landau, and Robin Stark for assistance in the field; Ronald Pursell for identifications of Fissidens and Richard Zander for critical determinations in the Pottiaceae; Jesus Munoz for information on the distribution of Grimmia americana; the granting of collecting permits from the U.S. Bureau of Land Management, Las Vegas office (Gay- le Marrs-Smith), Lake Mead National Recreation Area (Elizabeth Powell and Jennifer Haley), U.S.D.A. Forest Service (Kerwin Dewberry), Nevada State Parks, and the Nevada Test Site; Harold Robinson and Robert Ireland for reviewing the manuscript; and Bruce Allen, Bruce Lund, and Philip Medica for logistic support. LITERATURE CITED ANDERSON, L. E., H. A. CRUM, AND W. R. Buck. 1990. List of the mosses of North America north of Mexico. Bryologist 93:448—499. 2002] CANO, M. J., J. GUERRA, AND R. M. Ros. 1992. Crossidium seriatum (Pottiaceae, Musci) new to Europe. Bryol- ogist 95:280—282. Crum, H. A. 1994. Grimmia. Pp. 386—408 in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mexico. Pt. 1. Memoirs of the New York Botanical Garden 69, New York. AND L. E. ANDERSON. 1981. Mosses of Eastern North America, 2 vols. Columbia University Press, New York. AND W. R. Buck. 1994. Leskeaceae. Pp. 847-860 in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mexico. Pt. 2. Memoirs of the New York Botanical Garden 69, New York. Evans, A. W. 1917. 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Eckel (eds.), The moss flora of Mexico. Pt. 1. Memoirs of the New York Botanical Garden 69, New York. SCHOFIELD, W. B. AND H. A. Crum. 1972. Disjunctions in bryophytes. Annals of the Missouri Botanical Garden 59:174—202. SHABBARA, H. M. 1999. Three new records of Funariaceae from Egypt. Journal of Bryology 21:201—205. SMITH, D. K. 1994. Funariaceae. Pp. 427—442 in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mexico. Pt. 1. Memoirs of the New York Botanical Garden 69, New York. SPENCE, J. R. 1987. Pseudocrossidium aureum (Bartr.) Zand. (Pottiaceae, Musci) new to Utah. Great Basin Naturalist 47:347—348. . 1988. Checklist of the mosses of the Intermoun- tain West, USA. Great Basin Naturalist 48:394—401. STARK, L. R. 1997. Phenology and reproductive biology of Syntrichia inermis (Bryopsida, Pottiaceae) in the Mojave Desert. Bryologist 100:13—27. , B. D. MISHLER, AND D. N. McCLETCHIE. 1998. Sex expression and growth rates in natural populations of the desert soil crustal moss Syntrichia caninervis. Journal of Arid Environments 40:401—416. AND A. T. WHITTEMORE. 1992. Additions to the bryoflora of southern California. Bryologist 95:65— 67. AND A. T. WHITTEMORE. 2000. Bryophytes from the northern Mojave Desert. Southwestern Naturalist 45:226—232. STOTLER, R. AND B. CRANDALL-STOTLER. 1977. A checklist of the liverworts and hornworts of North America. Bryologist 80:405—428. WHITTEMORE, A. T. 1996. The taxonomy of Targionia (Targioniaceae) in North America. American Journal of Botany 83(6 suppl.):22—23. [Abstract.] ZANDER, R. H. 1977. Crossidium seriatum found in the U.S.A. Bryologist 80:170—171. . 1993. Genera of the Pottiaceae: Mosses of Harsh Environments. Bulletin of the Buffalo Society of Nat- ural Sciences Vol. 32. Buffalo, NY. . 1994a. Barbula. Pp. 286—296 in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mex- ico. Pt. 1. Memoirs of the New York Botanical Gar- den 69, New York. . 1994b. Pseudocrossidium. Pp. 296-299 in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mexico. Pt. 1. Memoirs of the New York Botanical Garden 69, New York. . 1994c. Weissia. Pp. 213-225 in A. J. Sharp, H. Crum, and P. M. Eckel (eds.), The moss flora of Mex- ico. Pt. 1. Memoirs of the New York Botanical Gar- den 69, New York. , L. R. STARK, AND G. MARRS-SMITH. 1995. Didy- modon nevadensis, a new species for North America, with comments on phenology. Bryologist 98:590— 595. MADRONO, Vol. 49, No. 1, pp. 54-58, 2002 NOTEWORTHY COLLECTIONS CALIFORNIA BACCHARIS MALIBUENSIS R. M. Beauch. & Henr. (AS- TERACEAE).—Orange Co., [Santa Ana Mountains] North Ranch Policy Plan Area [proposed for inclusion in the NCCP], Fremont Canyon, N. of Santiago Creek and immediately S. of major stream fork [of the Fremont Can- yon drainage], 33.7907°N 117.6801°W, UTM Zone 11 N3740484 E435012, ca. 305 m, 23 Aug 2000, Riefner & Wolf 20-732 (RSA). Previous knowledge. Baccharis malibuensis was de- scribed by Beauchamp and Henrickson in 1996 (Aliso 14: 197-203) as a narrow endemic restricted to the Malibu Creek drainage in the Santa Monica Mountains, extreme western Los Angeles County. At that time, the species was known from 5 localities on private land within an area just over 18 km’, growing on volcanic and sedimentary substrates in chaparral, openings in scrub, and in the un- derstory of Quercus agrifolia Née (Beauchamp & Hen- rickson 1996, loc. Cit.). Significance. The Riefner & Wolf collection of B. mal- ibuensis from the Santa Ana Mountains represents the first record for that range, the first record for Orange County, and a disjunction of 93 km south from the southernmost station in the Santa Monica Mountains. At the Fremont Canyon site, the plant was rare, growing at the base of a north-facing slope in the understory of a Q. agrifolia ri- parian woodland along an intermittent stream course. As- sociated species reported for the site include Symphori- carpos mollis Nutt., Heteromeles arbutifolia (Lindley) Roem., Rhamnus sp., Toxicodendron diversilobum (Torrey & A. Gray) E. Greene, Artemisia douglasiana Besser, Pip- tatherum milaceum (L.) Cosson, and Ambrosia psilostach- ya DC. This region of Fremont Canyon is primarily un- derlain by marine sandstones of the Williams Formation, which consists of very resistant, cliff-forming, white to brownish-gray feldspathic sandstone, pebbly sandstone, and conglomeratic sandstone (Morton 1999, Open-File Report 99-172, U.S. Geological Survey). Aside from the Palos Verdes headlands, the Santa Ana Mountains are the closest near-coastal range south of the Santa Monica Mountains. Two other taxa, Dudleya cy- mosa (Lem.) Britton & Rose ssp. ovatifolia (Britton) Mor- an and Nolina cismontana Dice, exhibit a similar pattern of disjunction between the Santa Monica and Santa Ana Mountains. It is not completely surprising, therefore, to find that B. malibuensis is present in both ranges. In the Santa Monica Mountains, B. malibuensis is known only from private lands and the documented occurrences are represented by populations of very small size, quite vul- nerable to extirpation by development (Beauchamp & Henrickson 1996, Joc. cit.). For this reason the species was added to list 1B of the California Native Plant Society’s inventory of rare and endangered vascular plants, with rarity-endangerment-distribution (RED) of 3-3-3, their highest threat rating (www.cnps.org/rareplants/inventory/ 6thEdition.htm). At the present time it appears that B. mal- ibuensis is also quite rare in the Santa Ana Mountains; only a single, pistillate, multi-branched subshrub approx- imately 8 dm high and 15 dm wide was located in 2000. Owing to the extremely rugged topography and relatively undisturbed character of the vegetation of this region it is likely that additional plants could be discovered in other nearby canyons that were not explored in 2000. Since B. malibuensis is also known from west- and south-facing slopes in clearings and dense chaparral in the Santa Mon- ica Mountains (Beauchamp & Henrickson 1996, loc. cit.) further surveys for B. malibuensis in similar habitats of the northern Santa Ana Mountains are warranted. —STEVE Boyb, Herbarium, Rancho Santa Ana Botanic Garden, 1500 N. College Avenue, Claremont, CA 91711. COLORADO AND NEw MEXIco ERIGERON OCHROLEUCUS Nutt. var. SCRIBNERI (Canby ex Rydb.) Cronquist (ASTERACEAE).—COLORADO: Weld County, rocky ridge 5.6 km N of Rockport, elev. 1830 m, TIIN R66W sect. 6 E% NE%, 16 May 2000, Dorn 8222 (COLO, RM). Previous knowledge. A range extension of 68 km from Albany County, Wyoming. Significance. First report for Colorado. SALIX ARIZONICA Dorn (SALICACEAE).—COLORA- DO: Conejos County, wet meadow along streamlet off Red Lake Trail Road 0.8 km from Hy. 17, elev. 3140 m, 37°04.7'N, 106°24.1'W, 6 July 2001, Dorn 8852 (COLO, MO, RM). Previous knowledge. A range extension of 105 km from Rio Arriba and Taos counties, New Mexico. Significance. First report for Colorado. SALIX DISCOLOR Muhl. (SALICACEAE).—COLORA- DO: Larimer County, bank of South Branch Boxelder Creek at County Road 37, elev. 2195 m, 40°57.7'N, 105°14.8’W, 29 May 2001, Dorn 8752, same plant 10 July 2001, Dorn 8894 (COLO, MO, RM). Previous knowledge. A range extension of 50 km from Laramie County, Wyoming. Significance. First report for Colorado. SALIX WOLFI Bebb var. WOLFII (SALICACEAE).—NEW MEXICO: Rio Arriba County, meadow along Osier Creek ca. 1.2 km SW of confluence with Rio de los Pinos, elev. 2925 m, 36°59.6'N, 106°20.6’W, 5 July 2001, Dorn 8847 (MO, NMC, RM, UNM). Previous knowledge. A range extension of ca. 40 m from adjacent Conejos County, Colorado or 2 km from nearest collection site in same county. Significance. First report for New Mexico. —RoBERT D. Dorn, Box 1471, Cheyenne, WY 82003. IDAHO CRYPTOGRAMMA STELLERI (S.G. Gmelin) Prantl (PTERI- DACEAE)—Boundary Co., Upper Priest Falls on Priest River, 1 km south of British Columbia border, 48°59’N, 116°55'W, rare on seepy, crumbly calcareous rock around waterfall in forest of Tsuga heterophylla and Thuja pli- cata, with Asplenium viride, ca. 950—1050 m, 20 Jul 2001, T. Spribille 11177 (ID). 2002] Previous knowledge. A widespread North American— Asiatic fern known from widely scattered localities in the western United States (E.R. Alverson, 1993, Cryptogram- ma. In: Flora of North America 2: 137—139) and infre- quent in adjacent British Columbia (G.W. Douglas et al., 1991, Vascular Plants of British Columbia). Significance. First report for Idaho, a range extension of approximately 330 km west from the nearest known locality on the east side of Glacier National Park, Glacier Co., Montana. VIOLA SELKIRKII Pursh ex Goldie (VIOLACEAE)—Bon- ner Co., upper end of Priest Lake, along Ruby Creek, ca. 48°50’N, 116°55’W; frequent in riparian Thuja plicata for- est, with Tiarella trifoliata, Viola glabella and Oplopanax horridus, ca. 730 m elev., 20 Jul 2001, T. Spribille & R. Merkel 11123 (ID). Previous knowledge. A widespread circumboreal spe- cies, rare in western North America, where it is known from widely scattered localities in Alaska, British Colum- bia, Alberta, Colorado (E. Hultén, 1968, Flora of Alaska and Neighboring Territories), New Mexico (W.C. Martin & C.R. Hutchins, 1980, Flora of New Mexico) and Mon- tana (T. Spribille et al., 2002, Noteworthy Collections, Madronio 49:55—58). Significance. First report for Idaho, a range extension of approximately 150 km west from the nearest known locality in the Whitefish Range, Lincoln Co., Montana. —Tosy SPRIBILLE, Kootenai National Forest, Fortine Ranger District, PO. Box 116, Fortine, MT 59918 (current address: Herbarium, Department of Systematic Botany, Albrecht von Haller Institute of Plant Sciences, University of Gottingen, Untere Karspiile 2, D-37073 Gottingen, Ger- many; e-mail toby.spribille@ gmx.de). MONTANA ALNUS RUBRA Bong. (BETULACEAE)—Lincoln Co., western Cabinet Mountains, Callahan Creek drainage, along Callahan Creek and North Callahan Creek at Mon- tana/Idaho state line and up to 3 km east into Montana, 48°26'30"N 115°57'—116°00'W, in alluvial bottoms and on moist slopes with Thuja plicata, Tsuga heterophylla and Betula papyrifera, 850-915 m, 16 Jun 1998, T. Spribille & M. Arvidson 7788 (USFS Fortine District Herbarium, COLO, MONTU, MRC). Previous knowledge. This primarily coastal species has a limited distribution in the interior and is usually asso- ciated with inland rainforest communities (Johnson & Steele, 1978, Northwest Sci. 52(3): 205—211; C.C. Lorain, 1988, Floristic history and distribution of coastal disjunct plants of the northern Rocky Mountains, M.Sc. thesis, Univ. Idaho). It is widespread along the Pacific Coast and is known from disjunct inland populations in British Co- lumbia, Washington and Idaho. Significance. First report for Montana, representing the eastern limits of species distribution, a contiguous range extension across the state line from nearby Boundary Co., Idaho. As in the Idaho locations, there appears to be in- trogression with Alnus incana ssp. tenuifolia in at least some individuals. The first reports of red alder from this area came from forest stand examination contractors who reported A/nus in the area as large as 59 cm dbh. AZOLLA MEXICANA Presl. (AZOLLACEAE)—Ravalli Co., McCalla Creek, app. 2 km west of Stevensville, NOTEWORTHY COLLECTIONS 55 46°30'20"N_ 114°07'40"W, locally common and forming small mats, with Callitriche heterophylla and Elodea can- adensis, 1030 m, 18 Sep 1999, W.E. Albert 3261 (MRC). Verified by PE Stickney (MRC). Previous knowledge. Widespread across the western states and British Columbia to South America, scattered east to the Mississippi River (T. A. Lumpkin, 1993, Azol- laceae. In: Flora of North America 2: 338-342). Significance. First report for Montana, probably intro- duced with recent disturbance, persisting and spreading in similar nearby drainages and a nearby slough. The nearest known collection stations are 320 km southwest and southeast, respectively, in Ada and Bannock Cos., Idaho (specimens at ID). BOTRYCHIUM PEDUNCULOSUM W.H. Wagner (OPHIO- GLOSSACEAE)—Lincoln Co., Big Creek, 48°44'30"N 115°28’30W, rare in moonwort genus communities on floodplains under Thuja plicata, 975 m, 24 Jul 1996, J. Vanderhorst 5609 (MONTU); Big Creek, 48°46'’N 115°27'W, 9 Aug 1997, J. Vanderhorst 5617 (MONTU); Cedar Creek, 10 km west of Libby, 48°24’35’N 115°40'59’W, with Thuja plicata, 1070 m, 1 Aug 1997, R. Ferriel s.n. (USFS Kootenai NF Herbarium); Quartz Creek, 17 km northwest of Libby, 48°30'50’N 115°42'07’W, with Tsuga heterophylla, Thuja plicata and Athyrium felix-femina, 910 m, 15 Aug 1997, R. Ferriel s.n. (USFS Kootenai NF Herbarium); Davis Creek, 20 km southwest of Trego, 48°31'37"N 114°57'40’W, in roadbed and powerline corridor, 1150 m, 22 Jul 1998, R. Ferriel RF98017 (USFS Kootenai NF Herbarium): Sanders Co., 0.8 km northeast of Noxon Rapids Dam, 47°57'53"N 115°43'21”W, mesic meadow, 735 m, 27 Jun 1999, R. Fer- riel RF99014 (USFS Kootenai NF Herbarium). Vander- horst 5609, 5617 determined by W. H. Wagner (MICH). Previous knowledge. A rare species of northwestern North America from southwestern Saskatchewan west to British Columbia and Oregon, east of the Coast/Cascade Ranges (W.H. Wagner & ES. Wagner, 1993, Ophioglos- saceae. In: Flora of North America 2: 85—106). Significance. First reports for Montana, a range exten- sion of approximately 130 km east from Pend Oreille Co., Washington. CAREX CHALCIOLEPIS Holm (CYPERACEAE)—Ravalli Co., Anaconda-Pintler Wilderness area, wet meadow that parallels both sides of the creek draining Hidden Lake, 45°55'10’N 113°33'00'W, 2500 m, Mooers & Mooers 998 (MONT). Determined by T. Spribille, verified by D.E Murray (ALA). Previous knowledge. A species of subalpine to alpine meadows in the southern and central Rocky Mountains, described by Holm (1903, Amer. J. Sci. 16: 17—44). Mur- ray (1969, Brittonia 21: 55—76) expressed doubt as to the occurrence of this species in Montana, because the ma- terial available to him was too immature to be certain of its identity. Significance. First report for Montana, representing the northern limits of species distribution, a range extension of over 300 km northwest from the nearest reported lo- cation in Park County, Wyoming. CAREX DEFLEXA Hornem. var. BooTTI L.H. Bailey (CY- PERACEAE)—Beaverhead Co., Beaverhead National Forest, Stine Mtn., West Pioneer Range, growing in Larix lyallii grove, 2640 m, 21 Jul 1968, S.F. Arno 29 (MON- TU); Beaverhead Co., Lima Peaks, one mile east of Gar- field Mtn., 8 miles south of Lima, common forming small patches of turf in stony, quartzite-derived soil of an alpine 56 MADRONO fellfield, 3030 m, 27 Jul 1989, P. Lesica & S. Cooper 4966 (MONTU); Missoula Co., Flathead National Forest, Lindy Peak, Mission Range, beneath stunted Larix lyallii, 2515 m, | Sep 1968, S.F. Arno 285 (MONTU); Missoula Co., Bitterroot Mtns., fellfield on Onehorse Ridge, Lolo Peak massif, 2575 m, 21 Jul 1971, K. Lackschewitz 2953 (MONTU); Ravalli Co., East Boulder Peak, growing in talus enclosure within alpine larch [stand], SE slope, 2710 m, 7 Aug 1968, K. Lackschewitz & T. Fageraas 596 (MONTU); Ravalli Co., dry, rocky crags, W-slope of the Castle Crags, 2590 m, 18 Aug 1970, K. Lackschewitz & Stuart 2365 (MONTU); Ravalli Co., NE slope of Canyon Peak, Canyon Lakes Basin, beneath alpine larch, 2590 m, 22 Aug 1971, K. Lackschewitz 3337 (MONTU); Ravalli Co., St. Joseph Peak, wind-timber zone, 2740 m, 24 Jul 1971, K. Lackschewitz & Gouaux 2979 (MONTU). All specimens determined by T. Spribille and verified by A.A. Reznicek (MICH). Previous knowledge. Although previously reported for Montana by Rydberg (1900, Flora of the Rocky Moun- tains) and EJ. Hermann (1970, Manual of the Carices of the Rocky Mountains and Colorado Basin, USDA Agr. Handb. 374) as Carex brevipes W. Boott, this species was subsumed under Carex rossii Boott in Hook. by C.L. Hitchcock et al. (1969, Vascular plants of the Pacific Northwest, Vol. 1), and has since escaped mention in the floras of Montana. The species is distinct from C. rossii both morphologically and ecologically, being a distinctly subalpine to alpine taxon with an apparent affinity for La- rix lyallii stands near the alpine timberline, in contrast to the more montane, xerothermic C. rossii. Furthermore, C. deflexa is more widespread across boreal North America than the primarily western C. rossi. Significance. These reports reaffirm the presence of this species in Montana. CAREX LACUSTRIS Willd. (CYPERACEAE)—Lake Co., small glacial pothole marsh ca. 6 km south of Swan Lake, 47°52'15"N 113°49'50"W, with Carex utriculata and C. lasiocarpa, 945 m, 15 Jul 1989, P. Lesica 4893 (MICH, MONTU); Swan River Valley, Lost Creek Fen, ca. 5 km south of village of Swan Lake, 47°52'55"N 113°49'42"W, in mossy saturated peat of fen, with Betula glandulosa and Carex lasiocarpa, 965 m, 19 Jun 1992, J.S. Shelly & S. Chadde 1652 (MICH). Both specimens determined by A.A. Reznicek (MICH). Previous knowledge. A species of the Great Lakes and Great Plains, rarely as far west as Idaho (M.L. Fernald, 1942, Rhodora 44: 281—331; Great Plains Flora Committee, 1977, Atlas of the Flora of the Great Plains), north-central Alberta (J.G. Packer, 1983, Flora of Alberta), and reportedly also British Columbia, although this report was based on a mis- identification (A. Ceska personal communication). Significance. First reports of this species for Montana. It joins a suite of species of the eastern deciduous wood- land region (e.g., Carex comosa, Carex pallescens, Dryop- teris cristata, Primula mistassinica) represented by dis- junct localities in northwestern Montana, northern Idaho and southeastern British Columbia. CAREX PALLESCENS L. (CYPERACEAE)—Ravalli Co., Bitterroot Valley along Bass Creek, 46°34'35"N 114°09'11”W, uncommon in moist meadow bordering Pi- nus ponderosa and Pseudotsuga menziesii with Juncus balticus, 1020 m, 6 Jul 1997, W. E. Albert 3167 (MICH); Bitterroot Valley, app. 18.4 km south and 2.4 km west of Florence, 46°28'40"N 114°09’'11”W, uncommon in season- ally-saturated meadow along a small stream with Populus trichocarpa, Pinus ponderosa, Carex illota and C. lanu- [Vol. 49 ginosa, 1085 m, 11 Jun 2000, W. E. Albert & B. Heidel s.n. (MONTU); Bitterroot Valley, south of Hamilton at Cory Place, 46°12’05’N 114°10’05’W, along waterway, 1090 m, 27 Jun 1979, J. Cory 1917 (MONTU). Albert 3167 determined by B. Heidel, verified by A.A. Reznicek (MICH); Cory 1917 determined as Carex torreyi by K. Lackschewitz, annotated to C. pallescens by A.A. Rezni- cek (MICH). Previous knowledge. An eastern species, introduced on Hornby Island, British Columbia from Europe or eastern North America (G.W. Douglas et al. 1994. Vascular Plants of British Columbia, Part 4), and recently discovered in Stevens Co., Washington, until recently not otherwise known from west of Great Lakes region. Significance. First reports for Montana, a range exten- sion of approximately 335 km southeast of Stevens Co., Washington (Bjork 3463, WS, ID). It is otherwise disjunct approximately 1700 km west from Duluth, Minnesota. CAREX PRAIREA Dewey (CYPERACEAE)—Flathead Co., northern Salish Mountains, confluence of Lime and Magnesia Creek drainages, approximately 6.4 km south of Trego, 48°38'30"N 114°52'30’W, locally common in matted clumps over 8—10 acres of bog birch fen complex with Potentilla fruticosa and Carex capillaris, 1060 m, 25 Jul 1995, F.J. Triepke 24 (USFS Fortine District Herbar- ium); Flathead Co., northern Salish Mountains, Magnesia Creek drainage, Magnesia Fen, 48°37'30"N 114°52'30’W, very common and abundant in fen, with Betula glandu- losa, Carex leptalea and Tomentypnum nitens, 1145 m, 21 Jun 1995, 7. Spribille 3355 (MICH), T. Spribille 3358 (USFS Fortine District Herbarium); same location, 1 Aug 1995, T. Spribille 3902 (COLO); Flathead Co., northern Salish Mountains, Blessed Creek ca. 0.75 km above con- fluence with Sunday Creek, 48°49'40’N 114°32'20’W, in rich calcareous fen, local in small patches, 1325 m, T. Spribille 7749 (COLO). T. Spribille 3355 verified by A.A. Reznicek (MICH). Previous knowledge. A widespread species of the boreal forest, Carex prairea has been reported from the western cordillera in Idaho (R. Davis, 1952, Flora of Idaho) and Wyoming (E. Hultén & M. Fries, 1986, Atlas of North European Vascular Plants North of the Tropic of Cancer, 3 vol.), although we have been unable to locate vouchers for these reports. In western Canada it is known from several sites in the Cariboo-Chilcotin region of central British Columbia (A. Roberts, 1983, A Field Guide to the Sedges of the Cariboo Forest Region, British Columbia. B.C. Min. For., Land Management Rep. No. 14; T-M.C. Taylor, 1980, The sedge family [Cyperaceae] of British Columbia, Royal B.C. Museum Handb. #43) and from central Alberta (J.G. Packer, 1983, Flora of Alberta). Significance. First reports for Montana, a range exten- sion of ca. 420 km south from the nearest known stations in west-central Alberta. CAREX VAGINATA Tausch (CYPERACEAE)—Lincoln Co., northern Salish Mountains, White Creek, just south of Forest Service Road 3529, 48°34’00"N, 114°56'30’W, 1200 m, in midmontane Picea glauca swamp with Rubus pubescens, and Carex disperma, 5 Jun 1996, T. Spribille & F.J. Triepke 5054 (USFS Fortine District Herbarium, | MICH); Lincoln Co., northern Salish Mountains, fen on White Creek along FS Rd. 36, 48°33'45’"N, 114°57'00"W, | 1125 m, 17 Jul 1998, T. Spribille & R.S. Wirt 7938 | (MONTU). Spribille & Triepke 5054 verified by A.A. Reznicek (MICH). Previous knowledge. A widespread pan-continental spe- ~ cies of the boreal forest from Alaska to Labrador (A.E. 2002] Porsild & W.J. Cody, 1980, Vascular Plants of Continental Northwest Territories, Canada, Natural Museum of Natu- ral Sciences, Ottawa), south to New York, Michigan and Minnesota. Significance. First reports for Montana and the western contiguous United States, a range extension of about 120 km south from the nearest location mapped by T.M.C. Taylor (The sedge family [Cyperaceae] of British Colum- bia, Royal B.C. Museum Handb. #43, 1980) in the south- ern Rocky Mountain Trench of British Columbia. CENTAURIUM ERYTHRAEA Rafin. (GENTIANACEAE)— Sanders Co., Cabinet Gorge Reservoir, road between Nox- on and Heron on south side of reservoir, ca. 5 km south of mouth of Elk Creek, locally common where powerline maintenance road clearing connects to the main road. 48°02'N 115°52'30"W, 670-730 m, 25 Aug 1997, T. Spri- bille 7429 (COLO). Verified by W.A. Weber (COLO). Previous knowledge. A Eurasian meadow species with medicinal uses, reported as established in northwestern North America from southern British Columbia (G.W. Douglas et al., 1990, Vascular plants of British Columbia, Part 2) south to California and inland to Idaho (C.L. Hitchcock et al. 1959, Vascular plants of the Pacific Northwest, Vol. 4). Significance. First report for Montana, representing the most inland station in western North America. ERIOGONUM VISHERI A. Nels. (POLYGONACEAE)— Carter Co., Powderville Road badlands, on the divide be- tween Dry Creek and Whitetail Creek, 45°46'18"N 104°55'42”, occasional on outcrops and outwash flats of Hell Creek Formation shale with Allium textile, Musineon divaricatum and Elymus lanceolatus, 948 m, 6 Jun 1997, B. Heidel 1540 (MONT); same location, 12 Jul 1997, J. Vanderhorst 5732 (MONT, MONTU). Previous knowledge. A regional endemic of the Great Plains, previously known only from North and South Da- kota (Great Plains Flora Committee, 1986, Flora of the Great Plains). Significance. First report for Montana, a range exten- sion of approximately 100 km southwest from Slope County, North Dakota and 150 km northwest from Har- ding Co., South Dakota. LESQUERELLA DOUGLASII S. Wats. (BRASSICACEAE)— Lincoln Co., Lake Koocanusa, Rexford Bench, 48°54’N 115°10'30"W, just west of Rexford along path; occurring infrequently and in small populations on sand in Pinus ponderosa/Purshia tridentata community, 775 m, 14 May 1999, F.J. Triepke & A. Stachurska 233 (COLO), same location, fruiting material, 26 Jun 2000, F.J. Triepke 246 (USFS Fortine District Herbarium). Triepke 246 verified by R. Hartman (RM). Previous knowledge. A Columbia Basin species of Washington, Oregon and British Columbia, known from a string of disjunct populations in the Rocky Mountain Trench of southeastern British Columbia. Rollins (1993, The Cruciferae of Continental North America) suggested it was to be expected in northwestern Montana. Significance. First report for Montana, a range exten- sion of ca. 40 km from the nearest known location along the Elk River near Grasmere, British Columbia (Spribille #760, UBC). MIMULUS RINGENS L. (SCROPHULARIACEAE)—Cho- teau Co., south shore of Missouri River, approximately 8 km east of Virgelle, 48°02'08”"N 110°09'06” W, seasonally flooded sandbar with Populus deltoides seedlings, Eleo- charis palustris and Helenium autumnale, 767 m, 7 Jul NOTEWORTHY COLLECTIONS 57 2000, B. Heidel 1952 (MONTU). Verified by R. Meinke (OSC). Previous knowledge. This primarily eastern species is known only from widely scattered western stations in Col- orado (W.A. Weber & R.C. Wittman, 1992, Catalog of the Colorado Flora: A Biodiversity Baseline), Idaho (R. Da- vis, 1952, Flora of Idaho), California (J.C. Hickman, ed. 1993. The Jepson Manual of the Higher Plants of Cali- fornia) and Washington (specimen at WTU). Significance. First report for Montana, a range exten- sion of at least 745 km west from Rolette Co., North Da- kota. RIBES LAXIFLORUM Pursh (GROSSULARIACEAE)— Lincoln Co., West Cabinet Range, south end of Little Spar Lake, 29 km SSW of Troy, 48°12'38”N 116°01'06’W, tall shrub field, 1675 m, 8 Sep 1997, E. Pederson 500 (MRC); same location, 26 Aug 1998, M. Arvidson & L. Ferguson 10I5 (MRC). Both specimens verified by PE Stickney (MRC). Previous knowledge. A Pacific coastal species, known inland from scattered stations in the Rocky Mountains of British Columbia (G.W. Douglas et al., 1990, Vascular Plants of British Columbia, Part 2), southwestern Alberta (C.L. Hitchcock & A. Cronquist, 1973, Flora of the Pacific Northwest), Colorado and New Mexico (A. Cronquist et al. 1997, Intermountain flora, Vol. 3, Part A.). Significance. First report for Montana, a range exten- sion of 13 km east from the nearest known location, at Halverson Creek, Bonner Co., Idaho, 1 km from the Mon- tana state line (Pederson 26, MRC). SENECIO CONGESTUS (R. Br.) DC. (ASTERACEAE)— Roosevelt Co., rangeland, no coll. date, received 11 Jun 1992, location information unavailable, Roosevelt County Extension Service s.n. (MONT). Determined by J. H. Ru- mely (MONT). Previous knowledge. A pan-continental boreal wetland species known from Newfoundland to Alaska and south from South Dakota and Iowa (Great Plains Flora Com- mittee 1986, Flora of the Great Plains) to Michigan (E.G. Voss, 1972—1996, Michigan Flora). Significance. First report for Montana, a range exten- sion of at least 30 km west from Divide Co., North Da- kota. VENTENATA DUBIA (Leers) Coss. & Dur. (POACEAE)— Ravalli Co., Skalkaho Creek, 46°09'57’N 113°55'54’W, common on dry roadsides with Poa compressa, Stipa co- mata. 1219 m, Jul 1995, W. E. Albert 3131 (MONT). Ver- ified by J. R. Rumely (MONT). Previous knowledge. A southern European species of dry grasslands introduced in western and northeastern North America, known from Idaho and Washington (C. L. Hitchcock et al. 1969. Vascular Plants of the Pacific Northwest, Vol. 1), southwestern British Columbia (G. W. Douglas et al., 1994, Vascular Plants of British Columbia, Part 4) and Utah (L. Allen & M. Curto, 1996, Madrono 43:337—338). Significance. First report for Montana, a range exten- sion of approximately 120 km east from Idaho Co., Idaho. VIOLA SELKIRKII Pursh ex Goldie (VIOLACEAE)—Lin- coln Co., Whitefish Range, Grave Creek, on south bank of creek ca. 2 km downstream of Williams Creek conflu- ence, 48°50’45"N_ 114°49’45”, in alluvial mixed forest of Betula papyrifera and conifers, with Aralia nudicaulis and Symphoricarpos albus; infrequent, only 30—40 plants found; 1030 m, 8 Jun 1999, T. Spribille & A. Stachurska 9081 (BHO). Verified by H. Ballard Jr. (BHO). 58 MADRONO Previous knowledge. This circumpolar boreal species is found in North America primarily in the eastern deciduous forests, but is also known only from widely scattered lo- calities in the Rocky Mountains south to Colorado (cf. E. Hultén, 1968, Flora of Alaska and Neighboring Territo- ries) and New Mexico (W.C. Martin & C.R. Hutchins, 1980, Flora of New Mexico). Significance. First report for Montana, a range exten- sion from southeastern British Columbia. —Tosy SPRIBILLE, Kootenai National Forest, Fortine Ranger District, RO. Box 116, Fortine, MT 59918 (current address: Herbarium, Department of Systematic Botany, Albrecht von Haller Institute of Plant Sciences, University of Gottingen, Untere Karsptile 2, D-37073 Gottingen, Ger- many; e-mail toby.spribille@gmx.de); BONNIE HEIDEL, Montana Natural Heritage Program, 1515 E 6th Ave., He- lena, MT 59620 (current address: Wyoming Natural Di- versity Database, University of Wyoming, P.O. Box 3381, Laramie, WY 82071-3381, e-mail bheidel@uwyo.edu); WALLACE E. ALBERT, 3653% Silverthorn Drive, Stevens- ville, MT 59870; E JAcK TRIEPKE, Kootenai National For- est, Fortine Ranger District, PO. Box 116, Fortine, MT 59918; Jim VANDERHORST, Natural Heritage Program, West Virginia Division of Natural Resources, P.O. Box 67, Ward Rd. Elkins, WV 26241-0067; and G. MICHAEL AR- VIDSON, Kootenai National Forest, Three Rivers Ranger District, 1437 Hwy 2 N, Troy, MT 59935. OREGON HIERACIUM CAESPITOSUM Dumort. (ASTERACEAE).— Wallowa Co., along Bear Creek Road, ca. 2 km S of Wal- lowa, with Dactylis glomerata, Phleum pratense, and [Vol. 49 Pseudotsuga menziesii. Also in pastures, along logging trails but absent in adjacent undisturbed forest, and along roads and riparian areas bordering Bear Creek, elev. 1090 m, T1S R42E sect. 3, Long. 117.55, Lat. 45.48, 23 July 2000, Brooks (OCS #197099); T1S R42E sect. 15, 17 July 2001, Dwire 1728 (OSC) (Verified by K. L. Chambers, OSC). Distribution extends south along the Bear Creek Trail into the Eagle Cap Wilderness, and north in the ri- parian areas bordering the Wallowa River. Previous knowledge. This species is also known as Hieracium pratense Tausch, an outdated synonym ap- pearing in Hitchcock and Cronquist (1973) and other western floras. Native to Eurasia, meadow hawkweed was probably introduced into the United States in the 1820's, and was first reported in the Pacific Northwest in Pend Orielle Co., Washington in 1969. It has become wide- spread throughout Washington, northern Idaho, and north- western Montana (Wilson et al. Rangelands 19:18—23, 1997; Toney et al., Northwest Science 72:198—209, 1998). It is spreading rapidly, primarily in montane meadows, pastures, and disturbed areas along roads and hillsides. Meadow hawkweed is a tenacious invader, and is listed as a noxious weed in Washington (Class B), Idaho, and Montana (Category 2). Significance. First report of the species for Oregon. Al- though present in the Bear Creek drainage, Wallowa Co. for perhaps 10 years, meadow hawkweed was only re- cently distinguished from native hawkweeds. An addition- al unvouchered population of Hieracium caespitosum has been reported from Hood River Co., Oregon. —KATHLEEN A. DwirE, Department of Forest Science, Oregon State University, Corvallis, OR 97331-5752, and CATHERINE G. PARKS, USDA Forest Service, Pacific Northwest Research Station, Grande, Oregon 97850. 1401 Gekeler Lane, La i MADRONO, Vol. 49, No. 1, p. 59, 2002 ERRATUM In Volume 48, No. 2, there was a typographical error in the title of the paper by Dieter Wilken (pages 116—122). The correct title should be A new Ipomopsis (Polemoniaceae) from the southwest USA and adjacent Mexico. 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TTI ES oie ca costes nates OE PR gh vino n 0 deg POD ED cs vesnestn sessions 115 Canopy MACROLICHENS FROM Four FoREST STANDS IN THE SOUTHERN SIERRA MIXED CONIFER FORESTS OF SEQUOIA/KINGS CANYON NATIONAL PARK David C. Shaw and St@RCHASNCKET ek Bo Soe. pe NL) ov nccc ese sescnccnsensonss 122 CALYSTEGIA SILVATICA (CONVULVULACEAE) IN WESTERN NORTH AMERICA R. K. Bromine... 4 a IE... eS he. 130 PNRUZIOIN oo e oo oo acc NT BF os ESN. cscs duacsnnccsececcaes 132 $ WASHINGTON 2 onc A ESS NS on SSE Sa ROSSA soa vecceesesesaccosess 132 WIGOREA a DF A cn WSS SSA osc cckeccccnneceesee 132 SEEING THINGS WHOLE: THE ESSENTIAL JOHN WESLEY POWELL, EDITED BY WILLIAM DEBuys Wall terran Te ed oon ic oo once cs RUNS Sev ose sesc con ecesweccevene 134 INVENTORY OF RARE AND ENDANGERED PLANTS OF CALIFORNIA, BY THE CALIFORNIA NATIVE PLANT Society, Davip P. TIBOR, CONVENING EDITOR ENTE OAD FIG 2 2) DD 12, Re me ets Oy |S A rn eee 135 PAO 7 US) SCHEDULE EON SPEARGRS 652i ccs coccscccean eceicacevadacccesavitaulecowvinvessdeuecwnndece 136 Maprono (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. Postmaster: Send address changes to MADRONO, Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. Editor—Dr. JOHN CALLAWAY Dept. of Environmental Science University of San Francisco 2130 Fulton Street San Francisco, CA 94117-1080 callaway @usfca.edu Book Editor—Jon E. 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Joun Litt_e, Sycamore Environmental Consultants, 6355 Riverside Blvd., Suite C, Sacramento, CA 95831; the Editor of MAprRONO; three elected Council Members: Bian Tan, Strybing Arboretum, Golden Gate Park, San Fran- cisco, CA 94122: James SHEvock, National Park Service, 1111 Jackson St., Suite 700, Oakland, CA 94607-4807. 510/ 817-1231; ANNE BRADLEY, USDA Forest Service, Pacific Southwest Region, 1323 Club Drive, Vallejo, CA 94592. abradley @fs.fed.us; Graduate Student Representative: KirstEN M. FisHerR, Jepson Herbarium, University of California, Berkeley, CA 94720. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). MApRONO, Vol. 49, No. 2, pp. 61-98, 2002 CATALOGUE OF NON-NATIVE VASCULAR PLANTS OCCURRING SPONTANEOUSLY IN CALIFORNIA BEYOND THOSE ADDRESSED IN THE JEPSON MANUAL—PART I FRED HRUSA California Department of Food and Agriculture, Plant Pest Diagnostics Center, 3294 Meadowview Rd., Sacramento, CA 95832-1448 FHrusa @cdfa.ca.gov BARBARA ERTTER University and Jepson Herbaria, University of California, Berkeley, CA 94720-2465 ANDREW SANDERS Botany and Plant Sciences Department, University of California, Riverside, CA 92521-0124 GORDON LEPPIG! Biological Sciences Department, Humboldt State University, Arcata, CA 95521 ELLEN DEAN UC Davis Herbarium, Section of Plant Biology, University of California, Davis, CA 95616 ABSTRACT A catalogue of 315 non-native vascular plant taxa documented as occurring spontaneously in California beyond those addressed in The Jepson Manual: Higher Plants of California is presented. The catalogue was compiled from new collections by the authors and others, previously existing herbarium specimens, formal publications, other printed reports, and direct communications with field botanists. Only reports backed by herbarium vouchers are accepted as adequately documented. Of the 315 species, 58 are fully or sparingly naturalized in relatively undisturbed wildland habitats, 53 are naturalized in disturbed areas, 34 are tenuously established or locally persisting, 13 are non-escaped weeds of greenhouse or similarly cultivated environments, 43 are presumed to be non-persisting casuals (waifs), for 110 there is no current information or observations available, and 4 have likely been extirpated. In addition, 13 reported taxa are here specifically excluded as based on erroneous information. Taxa highlighted as already being fully naturalized or potential pests are Amaranthus rudis, Brassica fruticulosa, Boehmeria cylindrica, Calys- tegia silvatica subsp. disjuncta, Cabomba caroliniana, Cotoneaster lacteus, Crataegus monogyna, Dit- trichia graveolens, Fumaria capreolata, Geranium purpureum, Geranium rotundifolium, Hedera canar- iensis, Limnobium laevigatum, Maytenus boaria, Pyracantha crenatoserrata, Salvinia molesta, Trifolium tomentosum, and Verbascum olympicum. Key Words: Weeds, non-native, invasive plants, pest plants, voucher specimens. The significance of invasive non-native plants has recently gained prominence, as evidence mounts for both the environmental and economic devastation such invasions can cause. A recent is- sue of BioScience (51[2] Feb. 2001), for example, is devoted to the topic of ““Global Movements of Invasive Plants and Fungi.”’ On the national level, various legislative initiatives have been proposed to address the problem, such as the Harmful Nonna- tive Weed Control Act (S. 198). At the local level, Weed Management Areas, established through the ' Present address: California Dept. of Fish and Game, Coastal Timberland Planning Program, 619 Second Street, Eureka, CA 95501. coordinated efforts of public and private agencies and organizations, now blanket most of California. Obviously, for all of these efforts to work prop- erly, accurate and comprehensive information needs to exist on which non-native plants occur within the area of concern and what potential level of threat they represent. While the average citizen might assume that this information is readily avail- able, especially in this age of electronic databases, the reality is unfortunately otherwise. This is pri- marily because, although a broad spectrum of pro- fessional biologists and amateur enthusiasts eagerly hunt down and keep track of rare native species, non-natives have been historically under-reported if not outright ignored. As a result, our existing knowledge of the identity, occurrence, frequency, 62 MADRONO and distribution of non-native plants is often sketchy or preliminary. California is by no means an exception to this rule, and the incomplete coverage of non-native species was one of the acknowledged short-com- ings of The Jepson Manual: Higher Plants of Cal- ifornia (Hickman 1993), which otherwise repre- sented the state-of-the-art coverage of California plants at the time of its publication. In a subsequent statistical analysis of the numbers and distribution of the non-native taxa reported in the Manual (Rej- manek and Randall 1994), mention was made of some clearly naturalized plants that were missing, but no compilation was attempted. As a precursor tally by Hrusa and Ertter, over 70 non-native plant taxa beyond those included in the Manual were not- ed as naturalized in California (Ertter 2000). More- over, the composition, frequency and distribution of plants in a given area is not static; this is particu- larly a feature of the non-native component, but even for native plants frequent updates to floristic accounts are necessary to maintain currentness (Yatskievych and Raveill 2001). The following cat- alogue is presented as the first installment of a con- tinuing comprehensive effort to follow through on this preliminary note by compiling existing reports and documenting new occurrences of plant taxa that occur spontaneously in California but which are not treated in The Jepson Manual. Such cata- loguing is intended to serve several functions: To help field botanists and consultants identify un- known species they come across during survey ac- tivities; to assist in the recognition of introduction pathways; and finally to provide further raw mate- rial for study of the invasion process itself. MATERIALS AND METHODS Data sources. Records of non-native vascular plants reported as growing without cultivation in California were reviewed and compiled. Data sources consisted of herbarium specimens, formal publications, other printed reports, and verbal com- munications with field botanists. Primary were our own and others’ plant collections; secondary were the numerous published local California floras and verbal reports provided to one or more of the au- thors. We have also included detailed records as available for most of those taxa mentioned by Rej- manek and Randall (1994) as absent from The Jep- son Manual. Because of the inevitable potential for misidentification, and following standard taxonom- ic practice, only reports that were backed up by hard documentation in the form of voucher speci- mens deposited in a publicly accessible herbarium are included in the current catalogue; this includes both those observed by one of the authors, stated in publication that such a voucher does exist, or verified by herbarium staff that one does exist. For example, the following taxa listed in Rejmanek and Randall (1994) are not included in Part I of the [Vol. 49 current catalog because testifying vouchers could not be located: Cordyline australis (Forst.f.) Endl. (Liliaceae s.l.), Cutandia memphitica (Spreng.) Richt (Poaceae) and Dodonaea viscosa Jacq. (Sap- indaceae). Of these taxa, a Cordyline has been ob- served by Hrusa at Salt Point, Sonoma Co., but a voucher confirming the specific application has not yet been acquired. The other two also may be es- tablished in California, but neither have as yet been so verified by the deposition of annotated speci- mens. Many of the printed or verbal reports were from federal and state agency botanists, native plant so- ciety members, weed control groups, and other un- published sources. Of particular importance were the plant samples submitted for identification by land managers, owners, farmers, ranchers, et al. to one or more herbaria. From these sources exact col- lection locality and ecological situation were some- times difficult to ascertain, and the material provid- ed was sometimes too poorly preserved to allow for more than an equivocal determination. The re- sult was that, until field confirmation or better col- lections can be obtained, some reports currently re- main unverified and are thus excluded from this compilation. Nevertheless, the value of the coop- eration of landowners, land managers, or other field people cannot be overstated. At the same time, the resultant catalogue is more than a simple report compilation. All reports were subject to verification by one or more authors, who also had the responsibility of determining natural- ization status. Correspondence with and the assis- tance of staff at herbaria worldwide were needed in the effort to track down documenting specimens for some taxa reported for California. Although one cannot absolutely prove the absence of a specimen, by a preponderance of evidence some putative re- cords were excluded, including several in published sources. For those taxa where a specimen was available, extensive identification efforts were often required to confirm identity, involving both world- wide taxonomic literature and herbarium speci- mens. In some particularly difficult cases, voucher specimens were sent to the appropriate specialist for identification or verification. Other problems, both taxonomic and nomenclatural, were resolved by electronic correspondence with the appropriate specialist, including several in Eurasia. Taxa re- ported for California, but for which documenting vouchers appear to be absent, and others deter- mined on examination to be misidentified or erro- neously reported, are listed and discussed in the RESULTS section. Scope of inclusion. Our decision of what to in- clude in the catalogue was based on three criteria: 1) The plant occurs spontaneously in California. ‘‘Spontaneous”’ as used here encompasses not only fully naturalized populations but any evidence of successful reproduction or initial appearance inde- 2002] pendent of targeted cultivation, including casuals, garden escapes, and weeds of greenhouse, nursery, garden, lawn, and agricultural field. This exception- ally broad definition was chosen to capture the pos- sible first appearance of potentially invasive spe- cies. However, long-persisting individuals that were intentionally planted, such as trees near abandoned homesites are not included unless they show evi- dence of successful reproduction, either sexual or vegetative; 2) The species was not addressed in The Jepson Manual, not even as an equivocal mention subordinate to another taxon. For example, Mentha spicata L. var. longifolia L. (Lamiaceae) is not in- cluded in the catalogue because it is mentioned in the Manual under M. spicata var. spicata, even though the entry is somewhat ambiguous and the variety is now fully documented as being natural- ized in California. Likewise, we do not report new regional occurrences for species already included in the Manual (e.g., plants reported only for northern California that are documented from the southern part of the state as well); 3) At least one reported locality is documented by a voucher specimen de- posited in a publicly accessible herbarium. Among the authors, at least fifteen vouchers were specifi- cally prepared for reports that had not previously been so documented. It should be recognized that a statement in print that a voucher was made does not guarantee that one was indeed deposited and it is possible that some taxa included in this catalogue will eventually, after further research, be found to be unverified. In addition, specimens proving oc- currence, but held in private collections were not included until a specimen or duplicates were made available for public scrutiny. Catalogue format. Individual taxon records in the catalogue are formatted as follows: TAXON NAME: Scientific name and author in accordance with the most recently available refer- ences (e.g., International Plant Names Index [http:// www.ipni.org/]; Flora of North America North of Mexico; Flora Europaea on-line [http://www.rbge. org.uk/forms/fe.html]; Catalogue of New World Grasses _ [http://mobot.mobot.org/W3T/Search/nwgc. html]). In cases of conflicting recent treatments, final decisions were based on our collective judgement. DISTRIBUTION (D): Reported occurrences in geographic subdivisions of California as described in The Jepson Manual, arranged alphabetically. CURRENT STATUS (CS): A brief description of current status as can best be determined. There is disagreement among invasion ecologists as to the scope of the term “‘naturalized’’ (Richardson et al. 2000). We follow the terminology of Richardson et al. but split their “‘naturalized’’ into subcategories: 1, “‘naturalized in wildlands’’, and 2, “‘naturalized (outside of wildlands)’’. It is recognized that dif- ferential occupation of these habitats is not mutu- ally exclusive, that assignment to one or the other category may be subjective, and that it is in addi- HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 63 tion a rare situation that, in California, has not been *“‘disturbed”” at some time in the past. We neverthe- less find the additional information provided in a classification distinguishing “‘naturalized in wild- lands’ and “naturalized (outside of wildlands)”’ potentially useful. This is because we feel that weedy species with a propensity to invade relative- ly natural or undisturbed areas are of different con- cern and may exhibit different ecological or repro- ductive strategies than are those plants which, al- though perhaps well-established, are restricted to agricultural conditions, roadsides or other heavily disturbed sites. The category ““NCI” (no current information) needs some explanation. In general these are plants confirmed to have occurred at one time in California, but for which no immediate in- formation is available as to their continued pres- ence, rate of spread, or naturalization. Many re- cords placed here could be assigned to another cat- egory based on the label data, but we preferred to maintain a conservative position on records that were often more than thirty years old, that is, these plants may be gone, or have spread widely during the ensuing years and we would be amiss in imply- ing more knowledge about their current status than we actually have. Naturalization categories are described and stan- dardized abbreviations, as used in the catalogue, are introduced and described in Table 1. DOCUMENTATION (DOC): Reports are ar- ranged alphabetically by county (in bold) as either a literature citation for previously published reports, or as specimen citation for newly documented oc- currences. Duplicate specimens might be housed in additional herbaria beyond those cited. Herbarium abbreviations are as used in /ndex Herbariorum (http://www.nybg.org/bsci/ih/ih.html). Generally literature cited as documentation includes specimen citations; however, collection or accession numbers, collectors names, dates and other details of location and habitat are only occasionally available in these sources. We expect that published floristic compi- lations maintain professional standards, and for published records where the specimen location is not specifically cited, the institution with which the author is or was affiliated houses the majority of the documented material. Known misapplications or misidentifications based on cited literature sourc- es alone should be brought to our attention. NOTES: Additional discussion or explanation as required or available, such as nomenclatural prob- lems, history of introduction or evidence of inva- siveness. In addition, for taxa believed to be be- coming common or rapidly expanding their ranges, a short comparison among similar species and/or identification clues are provided. RESULTS A total of 315 non-native vascular plant species meeting the criteria for inclusion are listed in Ap- 64 MADRONO [Vol. 49 TABLE 1. CURRENT STATUS CATEGORIES. Catalogue abbrev. Description NW Naturalized in wildlands: A population that appears to be successfully propagating (sexual- ly or vegetatively) and maintaining itself in a wildland situation. N Naturalized (outside of wildlands): A non-wildland population that is apparently of suffi- cient size or distribution to make the chances of its disappearance due to stochastic fluc- tuation negligible. Persistence tenuous: A wildland or non-wildland population that is of few enough individ- uals to make the chance of disappearance via stochastic fluctuation a possibility. C Casual: Synonymous with “‘waif”’ as used in The Jepson Manual to describe a spontane- ous occurrence that shows no evidence of successful propagation and spread. Greenhouse, nursery, garden weed: Non-escaped weed of greenhouse, nursery, garden, or No current information: Plants confirmed to have occurred at one time in California, but for which there is no immediate information available as to the continued presence, rate of spread, or naturalization. Many records placed here are assignable to another catego- ry based on the label data, but we prefer not to categorize these without more current GH/C other highly cultivated environment. NCI information. al Extirpated: Or reported as such. Habitat alteration or eradication programs in place have either eliminated these taxa from California or have nearly done so. pendix 1 and itemized in detail (Appendix 2). Both lists are organized as in The Jepson Manual, with Appendix 1 intended to facilitate the location of names in Appendix 2. The classification used in the catalogue to convey current naturalization status is summarized in Table 2. If a taxon had populations that were classifiable in different categories, the taxon as a whole was included in the category in- dicating more complete naturalization. For exam- ple, Salvinia molesta D.S. Mitch. (Salviniaceae) has several populations well-naturalized in both wild- land (NW) and non-wildland sites (N), plus is known several relatively small, localized popula- tions from private ponds (classed as TEN); it was thus classified as naturalized (NW) for the sum- mary Statistics. A subset of included species have the potential to achieve or have already achieved significant nat- uralized distributions or are pest plants, either as rapidly spreading invasive species (e.g., Dittrichia graveolens (L.) Greuter, Asteraceae) or as first Cal- ifornia appearances of well-known pest plants that were immediately targeted for eradication (e.g., Salvinia molesta). These were taxa rapidly expand- ing their former ranges, serious invaders in other TABLE 2. CURRENT STATUS SUMMARY. Explanation and definition of individual categories are provided in Table 1. Catalogue Definition abbrev. ‘Total Naturalized in wildlands NW 58 Naturalized (outside of wildlands) N 5)3} Tenuous/locally persisting TEN 34 Greenhouse, nursery, garden weed GH/C 13 Casual (Cc 43 No current information NCI 110 Extirpated EXT 4 parts of the world, plants already widespread and relatively commonly encountered, or plants appar- ently widespread but not heretofore correctly iden- tified [e.g., Hedera helix in part = Hedera canar- iensis Willd., (Araliaceae)]. The plants in this sub- set are listed in Table 3. They are provided a longer discussion and identification clues within their in- dividual Catalogue entries. Excluded taxa. It is important to note also the number of putative reports that were excluded from the current catalogue not from lack of documenta- tion, but because the reports were determined by the authors to be based on erroneous information. The importance of accurate identification of non- native naturalizing taxa cannot be overstated, there are numerous examples to show how control was only achieved once the proper identity of a pest was determined. A notable example is Salvinia molesta (Thomas and Room 1986); a similar, but currently unresolved example from California involves the genus Salsola (Chenopodiaceae) (M. Pitcairn per- sonal communication). Due to the taxonomic and nomenclatural complexities of name application to non-native taxa, the following list of thirteen (13) excluded reports is certainly not exhaustive: Achnatherum clandestinum (Hack.) Barkw. (Po- aceae): Misapplied in California to the similar A. brachychaetum (Godr.) Barkw. Plants from Mon- terey Co. sometimes identified as A. clandestinum have been recently redetermined by Barkworth (UTC) as A. brachychaetum. The identity of a spec- imen reported as this (Barkworth 1993), from San Luis Obispo Co., remains unverified, but a possible duplicate (or second collection from the same site by the same collector) at UC was annotated by Barkworth as A. caudatum (Trin.) Jacobs, a taxon highly similar to and perhaps conspecific with A. brachychaetum (see Vickery et al. 1986) suggesting 2002] HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 65 TABLE 3. POTENTIALLY SIGNIFICANT PLANTS. Fully naturalized species or first California appearances of well-known pest plants. Taxon Amaranthus rudis Brassica fruticulosa Boehmeria cylindrica Calystegia silvatica subsp. disjuncta Cabomba caroliniana Cotoneaster lacteus Crataegus monogyna Dittrichia graveolens Fumaria capreolata Geranium purpureum Geranium rotundifolium Hedera canariensis Limnobium laevigatum Maytenus boaria Pyracantha crenatoserrata Salvinia molesta Trifolium tomentosum Verbascum olympicum Summary of current plant behavior and status Well-established in widely separated localities. Rapidly spreading in mostly disturbed areas. Becoming common in the upper Sacramento River delta. Widespread in coastal areas, previously misidentified. Rapidly expanding, well-established aquatic invader. Dominant in some Sacramento River delta locations. Becoming locally common, expected more widely. Appearing in widely separated regions, often in stable habitats. Rapidly expanding. Unrecognized, probably more common than previously thought, likely expanding. Rapidly becoming more abundant. Widespread; overlooked previously, or rapidly expanding. Persistent invader of forest and woodland understories; previously con- fused with or included in H. helix. Rapid colonizer of open, still water, at low elevations. Can dominate to the exclusion of all other aquatic species. Aggressive invader of burns, forming long-lasting seedbanks. Widespread; invader of mesic ditchbanks, roadside depressions, open shorelines. Recent aquatic introduction into California where spreading in the Col- orado River and canal system. Widespread; previously overlooked or misidentified. Naturalized in wildland habitat, expanding for more than 20 years. that it may also be that taxon. Relationships among A. clandestinum, A. caudatum and A. brachychae- tum need critical review. Dichondra repens Forst. & Forst.f. (Convolvu- laceae): Misapplied in California. Acc. to Hortus III (1976) material cultivated as D. repens is actu- ally D. micrantha. See comments in the catalogue under the latter taxon. Glyceria declinata Bréb. (Poaceae): Acc. to Lep- pig, all specimens of putative Glyceria declinata from California are indistinguishable from G. oc- cidentalis. A discussion and treatment of variation in California Glyceria occidentalis is currently in preparation by Leppig. See also comments by Soza, Provance and Sanders (2000). Humulus americanus Nutt. sensu lato (Canna- baceae): The voucher for the report in Smith (1973, pg. 92), has not been located, but a new collection from the same locality (Echo Summit Lodge, El Dorado Co.) by Steve Matson has been verified by E. Small as H. lupulus var. neomexicanus A. Nelson & Cockerell. In Small’s treatment of Humulus in Flora of North America North of Mexico (3: 384— 387, 1997), H. americanus has been split among several varieties of H. lupulus native to North America: var. lupuloides E. Small, var. neomexi- canus, and var. pubescens E. Small. Although Small’s treatment reports var. neomexicanus from California, this apparent introduction is the only currently confirmed occurrence of the taxon in Cal- ifornia. We are rejecting the presence of H. amer- icanus S.\. because this name also applies to forms of H. lupulus (vars. pubescens and lupuloides) not in California. Hydrocharis morsus-ranae L. (Hydrocharita- ceae): Reports have all been misidentifications of, or misapplications to, Limnobium laevigatum. Both of these taxa are often called “‘frogbit’’, probably the source of the identity confusion. H. morsus-ran- dae is a possible threat to northern and high eleva- tion lakes in California, but has not yet been con- firmed in California outside of cultivation. Nothoscordum bivalve (L.) Britton (Liliaceae sensu lato): Apparently misapplied to Nothoscor- dum gracile (Ait.) Stearn s.l. (¢ncluding N. borbon- icum Kuntze = N. inodorum in the Jepson Manual). The cited location in Beauchamp (1986), “‘garden weed in Fallbrook”’ has a corresponding specimen at CDA of N. inodorum sensu auct. (=N. gracile or N. borbonicum). Passiflora mollissima (Kunth) L.H. Bailey (Pas- sifloraceae): Acc. to determinations by D. Goldman (BH) this name has been misapplied to P. tarmi- niana Coppens & Barney and P. mixta L.f., both of which are established in California. See d’Eckenbrugge et al. (2001). Polypogon elongatus Kunth (Poaceae): Misap- plied to P. imberbis (Phil.) Bjorkm. See catalogue for more information. Pueraria lobata (Willd.) Ohwi (Fabaceae): Ver- bal reports of kudzu vine in California have all been traced to native plants of Vitis californica. Pueraria has been cultivated in California as a cu- riosity, but in general, does not thrive under our dry 66 MADRONO [Vol. 49 TABLE 4. TIMING OF REPORTING FOR NEW CALIFORNIA RECORDS OF TAXA INCLUDED IN THE CURRENT CATALOGUE IN RELATION TO PUBLICATION OF THE JEPSON MANUAL. No. of taxa Relationship to The Jepson Manual 129 Discovered prior to the Manual but either not identified or a report not published. 110 Discovered (published report or not) subsequent to the Manual. 63 Discovered and a report published prior to the Manual. 11 Re-determination of pre-Manual species. 2 Accidentally excluded from the Manual, but the taxonomic treatment was completed and submitted prior to publication. Mediterranean climate. Locations discovered in northern Oregon in 2000 do apparently flower and fruit and are currently under pest evaluation. Salvia pratensis L. sensu stricto (Lamiaceae): Thomas C. Fuller collections from the region of Yreka (Siskiyou Co.) cited in Munz, PA. (1968, pg. 103) as S. pratensis have been redetermined at KW as Salvia virgata Jacq. See entry in the catalogue under that species for details. Sieglingia decumbens (L.) Bernh. (Poaceae): Misapplied in Matthews (1997) to Tribolium obli- terum (Hemzl.) Renvoize, first reported for North America in this catalogue. Cited also in Hitchcock and Chase (1950) as “escaped from cultivation, Berkeley”’, but no confirming evidence or speci- mens have been located. Spartina maritima (M.A. Curtis) Fern. (Po- aceae): Misapplied to Spartina anglica C.E. Hubb. For detailed explanation see Spicher and Josselyn (1985). Spartina spartinae (Trin.) Merr. (Poaceae): Mis- applied to S. densiflora Brongn. Reported from Humboldt Bay (Gerish 1979; Spicher and Josselyn 1985) and in Marin Co. at Greenwood Cove near Tiburon [Aug. 11, 1982, M.P. Wells s.n. (CAS, CDA)]. As a further complication, individual occurrence reports can be erroneous even if the plant itself is documented to occur spontaneously in California. As a particularly convoluted example, Amaranthus rudis (Amaranthaceae), which is included in the current catalogue on the basis of multiple docu- mented reports, had previously been cited in print (Munz 1974) only on the basis of a specimen from Pala in San Diego County, as A. tamariscinus Nutt. (misapplied to A. rudis). As it turns out, a duplicate of the voucher specimen (Townsend s.n., housed at CDA) has been re-determined as A. palmeri S. Wats. In other words, the report of A. rudis from near Pala in southern California is erroneous, even though other, mostly more recent reports substan- tiate the existence of this midwestern native in northern California. This example likewise under- scores the critical need to have available and ex- amine vouchers to confirm their identification. DISCUSSION Relation to the Jepson Manual. The Jepson Man- ual: Higher Plants of California (Hickman 1993) has for the past nine years provided California bot- anists with a new “‘standard”’ reference, supplant- ing the more than thirty-year old A California Flora (and its 1968 Supplement) by P.A. Munz and D.D. Keck. As earlier noted, non-native plants have his- torically been under-collected. Because floristic works such as the Manual used herbarium collec- tions as the fundamental scientific standard, the re- sult has been a general under-representation of non- native species. It is hoped the current paper, along with the burgeoning interest in non-native taxa as potential pest plants, will reverse this tendency. In addition to those species that were omitted from the Manual because of the lack of accessible herbarium documentation, others were not included for a variety of reasons. In particular, the Jepson Manual Project deliberately excluded ‘casual’ (‘waif’) species, unconfirmed naturalization re- ports, or material apparently persistent from culti- vation. While this was a reasonable decision for the Manual, we believe that even these marginal oc- currences are worth reporting, given that all taxa that are now naturalized and well-established com- ponents of the State flora began as rare casuals or as small populations. Finally, there are plants in the current catalogue that have appeared as spontane- ous occurrences in California subsequent to the clo- sure of treatments for the Manual. Worth mentioning in particular are those natu- ralized taxa that were reported in A California Flo- ra or adequately documented in the intervening pe- riod but which were not treated in the Manual. The reasons these taxa were excluded are diverse. Some had been included in A California Flora but current information as to their naturalization status was not available. For others, individual authors may have decided that the inclusion of marginally established or rarely encountered weeds unnecessarily compli- cated identification keys and/or in general did not benefit the majority of users. Again, while this ra- tionale may have been justified for the purposes of the Manual, this excluded group of plants is in- cluded in the current catalogue in order to assist those who have diverse interests concerning inva- Sive taxa. The temporal relationship of species in the cur- rent catalogue to The Jepson Manual is summa- rized in Table 4. 2002] Significance of new reports. The often pestifer- ous nature of the most prominent weeds in Cali- fornia is well-recognized by the layman and pro- fessional alike (e.g., Bossard et al. 2000) What is not so universally understood is that these wide- spread and/or noxious pests generally began their occupation of California as seemingly innocuous roadside waifs, occasional garden volunteers, minor seed source contaminants, or localized populations remaining for some time in other non-prominent situations. The California Dept. of Food and Agri- culture (CDFA) rates noxious weeds, and applies control efforts accordingly, not by their current af- fect on agricultural economics, but by their poten- tial effect, the position being that control is both most biologically and cost effective when the pop- ulations are small, before the plant has become a widespread, established pest. Data have been as- sembled and published in support of this position (Bayer 1999). Although CDFA is concerned pri- marily with weeds affecting agricultural endeavors, wildland pests are no different nor are these con- ditions mutually exclusive; it is important to ag- gressively control these organisms before they be- come too widespread for control to be effective or eradication possible. Addressing only the most widespread and well-established weeds in Califor- nia leaves a gap in our knowledge that may affect the future of California’s agricultural activities, pristine wildlands and outdoor recreation areas. That this is an important gap may be recognized when one considers that every presently wide- spread weed in California began long ago as a “‘ca- sual”? [Centaurea solstitialis L. (Asteraceae), Lep- idium latifolium L. (Brassicaceae)], or ““occasional garden escape” [Cyftisus scoparius (L.) Link, Gen- ista monspessulana (L.) L. Johnson (both Faba- Geae)p While it cannot be specifically claimed that any plant appearing in this catalogue will ultimately be- come more than even a local pest, the naturalization of non-native plants may ultimately have long-last- ing impacts. The truth of this statement is dramat- ically evident when perusing S. B. Parish’s 1920 synopsis of immigrant plants of southern Califor- nia, in the number of species that were merely pre- sent as scattered waifs in 1920 but which are now both fully naturalized and widespread. For exam- ple, Potamogeton crispus L. (Potamogetonaceae) was known in California from a single collection at Corona in 1918, while Pennisetum villosum R. Br. (Poaceae) was a “local fugitive from cultiva- tion” that Parish was aware of only from Santa Barbara and Ventura counties. Even currently ca- sual taxa have a potential long-term impact be- cause, unlike many kinds of environmental pollut- ants that are eventually neutralized within or dis- appear from the system upon termination of the source, biological pollutants such as naturalizing non-native plants are self-perpetuating, often in- creasing in prominence and distribution with time HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 67 regardless of whether the source has been termi- nated (O’Kennon et al. 1999). Moreover, as stated above, the cost of control rises linearly as popula- tion size increases (Bayer 1999; Rejmanek 2000); because populations often enlarge logarithmically, ultimate control costs for populations left uncon- trolled may exceed available resources. As a result, awareness of the identity and ecology of a potential or incipient weed population is not only important, it is also economically imperative—if control or eradication are eventual goals. We hope that this compilation will spur two ac- tivities: first, further exploration for non-native spontaneous plants in California; and second, clar- ification of the current status of the taxa on this list, especially those classified as “‘No Current Infor- mation (NCI)’”’. Clearly, the disturbed areas on which many of these latter taxa established in the earlier decades of this century have subsequently been stabilized by parking lots or other intensive development; however, other populations may have expanded their range, but are on private land no longer readily accessible. On-going need for vouchered reports. The pre- sent compilation is not exhaustive for several rea- sons. First and foremost is the rapid rate at which new weedy taxa are being found in a state as large and geographically diverse as California. Secondly, a compilation such as this, which includes many records known only from obscure locations or sin- gle sites in California, comprises a major effort by botanical collectors of modern and previous eras; without their past and ongoing efforts this collation would not exist. With this in mind, as California’s increase in human population and consequent plant introductions continues, it is important that the col- lection and documentation of non-native taxa, by the submission of specimens to public collections, continue as well. As a final note we wish to again emphasize the absolute need for specimen documentation of new reports of spontaneous non-native plants (see Dean and Hrusa 2000 for instructions on collecting and documenting plant occurrences). The fact that de- termining and verifying these species does not keep pace with the introduction rate is largely a reflec- tion of the difficulty in applying names to plants whose geographic origins are not known and which may be cultivated forms or which have become phenetically modified in their new habitats. For this reason it has been discouraging to realize how many seemingly authoritative reports, via both agencies and private organizations, lack confirming specimen material. Indeed, some lacked even the documentation of who applied the name and when, or where the plant was reported to grow. Without such information a reliable understanding of which taxon is actually represented is not possible. The ramifications of this information gap are great; eco- logical behavior, environmental tolerances, devel- 68 MADRONO opmental and phenological patterns are often spe- cies-specific, and control measures, whether phys- ical, chemical, or biological, may not be effective if the totality of life history is not correctly under- stood. The essential first step toward accessing whatever critical information is available is having an accurate identification (for a case study, see Thomas and Room 1986). The addition of so many non-native pest plants with the potential for becoming naturalized further underscores just how much the weed invasion of California should be of continuing economic and ecological concern. The current catalogue, while the result of intensive research in both the field and herbarium, is by no means complete; rather it serves to point out the extent to which non-native plants must be constantly sought out and reported. ACKNOWLEDGMENTS The current effort is dedicated to Thomas C. Fuller, Elizabeth McClintock, G. Douglas Barbe, and the late June McCaskill, our predecessors in the arena of docu- menting new occurrences of non-native plants in Califor- nia. Sincere appreciation and acknowledgment of our debt is also extended to the long list of individuals whose col- lective contributions, ranging from first collections of new occurrences to identifications of problematic taxa to other forms of assistance, made the catalogue possible: Carlos Aedo, Lowell Ahart, Mary Barkworth, James Barry, Steve Boyd, Robin Breckenridge, Richard Brummitt, Adolf Ces- ka, Steve Clemants, D. Gail DeLalla, Thomas Delendick, Joe DiTomaso, Ed Finley, Jesse Geissow, Jeff Glazner, D. Goldman, Jeff Hart, Evelyn Healy, Nancy Hillyard, Kevin Hintsa, Margaret Hurlbert, Lawrence Janeway, John Kar- tesz, David Keil, Dean Kelch, Rod Kerr, Dave Kratville, Elaine Mahaffey, Steve Matson, Mary Ann (Corky) Mat- thews, Dick Moe, Tony Morosco, Sergei Mosyakin, Barry Meyers-Rice, Robert Ornduff, Vern Oswald, Elizabeth Painter, David Philcox, Mike Pitcairn, Robert Preston, John Randall, Marcel Rejmanek, Tom Rosatti, Jake Ruygt, Mary DeValle Sanvictores, Rudi Schmid, Steve Schoenig, Teresa Sholars, Ernest Small, James P. Smith, John Stroth- er, Dean Wm. Taylor, Chris Thayer, Gordon Tucker, Mi- chael Vincent, Margriet Wetherwax, Dieter Wilken, Jo- seph Wipff, and Vern Yadon. Special thanks to these sharp-eyed individuals and any others, inadvertently over- looked, who have made vouchers or brought reports of new spontaneous non-natives to our attention. 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Flora of the Santa Cruz Mountains of California. Stanford University Press, Palo Alto, CA. THOMAS, P. A. AND P. M. Room. 1986. Taxonomy and control of Salvinia molesta. Nature: 320, 17 April. THURBER, G. 1880. Gramineae. Jn S. Watson, Botany of California, Vol. II. John Wilson & Son, University Press, Cambridge, MA. TWISSELMANN, E. 1956. Flora of the Temblor Range. The Wasmann Journal of Biology 14:161—300. TYLER, V. E. 1982. The new honest herbal; a sensible guide to herbs and related remedies. Stickley Co., Philadelphia, PA. USDA, NRCS. 2001. The PLANTS Database, Version 3.1 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. VICKERY, J. W., S. W. L. JACOBS, AND J. EVERETT. 1986. Taxonomic studies in Stipa (Poaceae) in Australia. Telopea 3:1—338. VINCENT, M. A. 1997. Noteworthy collections. Madrofo 44:305—306. WILKEN, D. 1993. Balsaminaceae. Addendum to the first and second printings of the Jepson manual. The Jep- son Globe 5:2. YADON, V. 1995. Checklist of the vascular plants of Mon- terey County, California. California Native Plant So- ciety, Sacramento, CA. YATSKIEVYCH, G. AND J. A. RAVEILL. 2001. Notes on the increasing proportion of non-native angiosperms in the Missouri flora, with reports of three new genera for the state. Sida 19:701—709. ZOHARY, M. AND D. HELLER. 1984. The genus Trifolium. The Israeli Academy of Sciences and Humanities, Je- rusalem, Israel. APPENDIX 1 LisT OF TAXA WITH NATURALIZATION CLASS List of non-native vascular plant taxa occurring spon- taneously in California but not included in The Jepson Manual. Organization as in the Manual. Current Status subcategories in parentheses: NW = naturalized in wild- lands, N = naturalized (outside of wildlands), TEN = per- sistence tenuous, C = casual (waif), GH/C = greenhouse/ cultivation, EXT = extirpated. See Table 1 and Appen- dix 2. FERNS AND ALLIES Salviniaceae Salvinia molesta D.S. Mitch., (NW). Selaginellaceae Selaginella kraussiana (Kunze) A. Braun, (N). Ephedraceae Ephedra altissima Desf., (NCI). Ephedra distachya L., (NW). CONIFERS Pinaceae Pinus halepensis P. Mill., (N). Pinus pinea L., (NW). ANGIOSPERMS: DICOTS Acanthaceae Acanthus mollis L., (C). [Vol. 49 Aceraceae Acer campestre L., (N). Acer saccharinum L., (NW). Aizoaceae Galenia pubescens (Ecklon & Zeyher) Druce var. pubes- cens, (N). Amaranthaceae Amaranthus blitum L. subsp. emarginatus (Uline & Bray) Carretero, (C). Amaranthus rudis Sauer, (N). Brayulinea densa (Willd.) Small, (NCI). Froelichia gracilis (Hook.) Mog., (NCI). Anacardiaceae Rhus lancea L.f., (N). Schinus polygamus (Cav.) Cabr., (N). Apocynaceae Nerium oleander L., (NW). Aquifoliaceae Ilex X attenuata Ashe, (NCI). Araliaceae Hedera canariensis Willd., (NW). Asclepiadaceae Asclepias fruticosa L., (C). Calotropis procera (Ait.) Ait. f., (C). Cynanchum louiseae Kartesz & Gandhi, (N). Asteraceae Achillea filipendulina Lam., (NCI). Artemisia vulgaris L. sensu stricto, (NCI). Aster novae-angliae L., (NCI). Carthamus oxyacantha M. Bieb., (C). Centaurea babylonica L., (C). Chrysanthemum balsamita L., (C). Cirsium scabrum (Poir.) Bonnett & Barratte, (NCI). Coreopsis grandiflora Hogg. ex Sweet, (C). Cotula mexicana (DC.) Cabrera, (N). Crepis tectorum L., (N). Dendranthema X grandiflorum Kitam., (NCI). Dittrichia graveolens (L.) Greuter, (NW). Emilia fosbergti D.H. Nicolson, (GH/C). Emilia sonchifolia (L.) DC. ex Wight, (GH/C). Grindelia papposa Nesom & Suh, (NCI). Helianthus tuberosus L., (C). Helichrysum petiolare Hilliard & B. L. Burtt., (NW). Heliomeris multiflora Nutt. multiflora, (TEN). Helipterum roseum (Hook.) Benth., (NCI). Lasiospermum bipinnatum (Thunb.) Druce, (NW). Leontodon muelleri (Schultz-Bip.) Fiori, (NCI). Oncosiphon piluliferum (L.f.) Kallersjo, (N). Osteospermum fruticosum (L.) Norl., (NW). Pericallis cruenta (DC. non Roth) Webb. & Berth., (NCI). Porophyllum ruderale (Jacq.) Cass., (C). Ratibida columnifera (Nutt.) Wooton & Standl., (NCI). Senecio squalidus L., (NCI. Tragopogon hybridus L., (NCI). Tripleurospermum maritimum (L.) W. D. J. Koch, (TEN). 2002] Balsaminaceae Impatiens balfouri J.D. Hook., (N). Impatiens noli-tangere L., (N). Berberidaceae Berberis darwinii Hook., (NCI). Betulaceae Alnus cordata (Loisel.) Duby, (C). Bignoniaceae Campsis radicans (L.) Bureau, (TEN). Catalpa bignonioides Walter, (NW). Boraginaceae Echium lusitanicum L., (TEN). Pentaglottis sempervirens (L.) Tausch ex Bailey, (NCI). Brassicaceae Brassica fruticulosa Cyrillo, (N). Cardamine flexuosa With., (N). Coincya monensis (L.) Greuter & Burdet, (NW). Iberis umbellata L., (C). Rorippa sylvestris (L.) Besser, (TEN). Cabombaceae Cabomba caroliniana A. Gray, (NW). Campanulaceae Campanula medium L., (C). Lobelia erinus L., (C). Caprifoliaceae Leycesteria formosa Wallich., (NCI). Viburnum tinus L., (NCI). Caryophyllaceae Silene pseudatocion Desf., (NCI). Celastraceae Maytenus boaria Molina, (NW). Chenopodiaceae Atriplex muelleri Benth., (NCI). Chenopodium watsonii A. Nels., (NCI). Salsola kali L. subsp. pontica (Pallas) Mosyakin, (NCI). Cistaceae Halimium lasianthum (Lam.) Spach, (NCI). Convolvulaceae Calystegia silvatica (Kit.) Griseb. subsp. disjuncta Brum- mitt, (N). Convolvulus tricolor L., (NCI). Dichondra micrantha Urb., (NCI). Ipomoea aquatica Forssk., (C). Ipomoea lacunosa L., (C). Ipomoea quamoclit L., (C). HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 71 Crassulaceae Crassula multicava Lem., (NW). Sedum album L., (N). Sedum dendroideum Sesse & Moc. ex DC., (NCI). Cucurbitaceae Cucumis anguria L., (NCI). Cucurbita ficifolia Bouche, (NCI). Cucurbita pepo L. var. medullosa Alef., (C). Cuscutaceae Cuscuta reflexa Roxb., (EXT). Droseraceae Drosera aliciae Hamet, (NCI). Drosera capensis L., (NCI). Drosera tracyi MacFarlane, (NW). Ebenaceae Diospyros virginiana L. var. virginiana, (NW). Elaeocarpaceae Aristotelia chilensis (Molina) Stuntz, (TEN). Muntingia calabura L., (GH/C). Escalloniaceae Escallonia macrantha Hook. & Arn., (NCI). Euphorbiaceae Euphorbia characias L., (NCI). Euphorbia cyathophora Murr., (NCI). Euphorbia dendroides L., (NW). Euphorbia heterophylla L., (NCI). Euphorbia hirta L., (N). Euphorbia hypericifolia L., (GH/C). Euphorbia marginata Pursh, (NCI). Euphorbia myrsinites L., (NCI. Euphorbia rigida M. Bieb., (NCI). Euphorbia terracina L., (NW). Sapium sebiferum (L.) Roxb., (NW). Fabaceae Astragalus cicer L., (TEN). Cassia nemophila A. Cunn., (TEN). Ceratonia siliqua L., (NW). Coronilla valentina L., (NW). Dolichos lignosus Pers., (NCI). Genista monosperma (L.) Lam, non Link, nec Del., (NW). Gleditsia triacanthos L., (NW). Lathyrus sativus L., (C). Ononis alopecuroides L., (NW). Robinia hispida L., (N). Senna artemisioides (Gaudich. ex DC.) Randell, (N). Senna obtusifolia (L.) H.S. Irwin & Barneby, (N). Sesbania punicea (Cav.) Benth, (N). Trifolium alexandrinum L., (NCI). Trifolium cernuum Brot., (N). Trifolium gemellum Poir. ex Willd., (N). Trifolium resupinatum L., (NCI). Trifolium retusum L., (NW). Trifolium stellatum L., (NCI). Trifolium striatum L., (NCI). Trifolium tomentosum Willk. ex Nyman, (NW). 72 Trifolium vesiculosum Savi, (N). Trigonella corniculata L., (NCI). Trigonella foenum-graecum L., (NCI). Vicia bithynica (L.) L., (NCI). Fagaceae Quercus ilex L., (N). Geraniaceae Geranium columbinum L., (NCI). Geranium lucidum L., (N). Geranium purpureum Vill., (NW). Geranium pyrenaicum Burm. f., (C). Geranium rotundifolium L., (NW). Geranium texanum (Trel.) A. Heller, (NCI). Hamamelidaceae Liquidambar styraciflua L., (TEN). Hydrophyllaceae Wigandia caracasana HBK., (NCI). Hypericaceae Hypericum androsaemum L., (N). Hypericum calycinum L., (TEN). Hypericum hookerianum Wight & Arn., (TEN). Lamiaceae Calamintha sylvatica Bromf. subsp. ascendens (JordaN). P.W. Ball, (TEN). Cedronella canariensis (L.) Willd. ex Webb & Berth., (NCI). Galeopsis tetrahit L., (NCI). Lamiastrum galeobdolon (L.) Ehrend. & Polatsch., (TEN). Lavandula stoechas L., (C). Mentha X villosa Huds., (NCI). Monarda citriodora Cerv., (N). Rosmarinus officinalis L., (C). Salvia longistyla Benth., (N). Salvia microphylla Benth., (NCI). Salvia reflexa Hornem., (GH/C). Salvia virgata Jacq., (NCI). Scutellaria caerulea M. & S., (C). Stachys floridana Shuttlew., (GH/C). Lauraceae Cinnamomum camphora (L.) J. Presl, (C). Laurus nobilis L., (TEN). Lentibulariaceae Utricularia subulata L., (NCI). Limnanthaceae Limnanthes macounii Trel., (N). Linaceae Linum trigynum L., (NCI). Malvaceae Anisodontea capensis (L.) Bates, (C). Anoda pentaschista A. Gray, (NCI). Gossypium hirsutum L., (NCI). MADRONO Hoheria populnea A. Cunn., (NCI). Lavatera olbia L., (NCI). Lavatera trimestris L., (NCI). Malva verticillata L., (NCI). Sida spinosa L., (NCI). Moraceae Fatoua villosa (Thunb.) Nakai, (GH/C). Ficus palmata Forssk., (NCI). Nymphaeaceae Nymphaea alba L., (NCI). Oleaceae Fraxinus uhdei (Wenz.) Lingel., (NW). Ligustrum lucidum W.T. Aiton, (NW). Ligustrum ovalifolium Hassk., (NW). Olea africana Mill., (TEN). Onagraceae Fuchsia magellanica Lam., (NW). Fuchsia X hybrida Voss., (NCI). Orobanchaceae Orobanche hederae Duby, (TEN). Papaveraceae Fumaria capreolata L., (N). Papaver X hybridum L., (NCI). Passifloraceae Passiflora caerulea L., (N). Passiflora manicata (Juss.) Pers., (NCI). Passiflora mixta L. f., (N). Passiflora tarminiana Coppens & Barney, (N). Pedaliaceae Sesamum indicum L., (C). Plumbaginaceae [Vol. 49 Limonium ramosissimum (Poir.) Maire subsp. provinciale (Pignatti) Pignatti, (NW). Polygalaceae Polygala myrtifolia L., (NCI). Polygonaceae Polygonum multiflorum Thunb., (GH/C). Polygonum orientale L., (C). Ranunculaceae Caltha palustris L., (NW). Clematis terniflora DC., (TEN). Clematis vitalba L., (NCI). Nigella damascena L., (N). Ranunculus cortusifolius L., (TEN). Rhamnaceae Ziziphus jujuba L., (C). 2002] HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 73) Rosaceae Cotoneaster lacteus W.W. Smith (NW). Crataegus monogyna Jacquin, (NW). Cydonia oblonga Mill., (NCI). Cydonia sinensis Thouin, (NCI). Eriobotrya japonica Lindl., (NCI). Filipendula vulgaris Moench, (NCI). Photinia davidsoniae Rehd. & Wilson, (NCI). Potentilla anglica Laicharding, (NW). Potentilla reptans L., (TEN). Prunus laurocerasus L., (NW). Prunus persica (L.) Batsch, (C). Prunus serrulata Lindl., (C). Pyracantha coccinea M. Roem., (TEN). Pyracantha crenatoserrata (Hance) Rehder, (N). Pyracantha crenulata (D. DON). M. Roem., (TEN). Pyrus communis L., (N). Rosa multiflora Thunb. ex Murray, (NCI). Rubus ulmifolius Schott var. ulmifolius, (N). Rubiaceae Coprosma repens A. Rich., (N). Salicaceae Populus nigra L. cv. “‘Ttalica”’, (TEN). Sarraceniaceae Sarracenia aff. rubra Walter, (N). Scrophulariaceae Anarrhinum bellidifolium (L.) Willd., (C). Limnophila X ludoviciana Thieret, (TEN). Mazus japonicus Kuntze, (GH/C). Penstemon strictus Benth., (TEN). Penstemon subglaber Rydb., (TEN). Scrophularia peregrina L., (N). Verbascum olympicum Boiss. non Bunyard, (NW). Solanaceae Atropa belladonna L., (NCI). Capsicum annuum L., (NCI). Cestrum parqui LU Her., (NCI). Lycium ferocissimum Meirs, (NCI). Nicotiana X sanderae Hort. ex Wats., (C). Nicotiana tabacum L., (C). Petunia violacea Lindl., (NCI). Solanum gayanum (Remy) Phil. f., (NCI). Solanum scabrum Mill., (C). Solanum villosum Mill., (C). Urticaceae Boehmeria cylindrica (L.) Sw., (NW). Laportea aestuans (L.) Chew, (GH/C). Verbenaceae Verbena rigida Spreng., (NCI). Vitex agnus-castus L., (NCI). Vitaceae Cissus antarctica Venten., (NCI). Vitis aestivalis Michx., (NCI). Vitis rupestris Scheele, (TEN). Zy gophyllaceae Peganum harmala L., (EXT). ANGIOSPERMS—MONOCOTS Alismataceae Sagittaria brevirostra Mackenzie & Bush, (NCI). Sagittaria rigida Pursh, (NW). Araceae Arum palestinum Boiss., (TEN). Dracunculus vulgaris Schott, (N). Pinellia ternata (Thunberg) Makino, (NCI). Cyperaceae Bulbostylis barbata Kunth, (GH/C). Cyperus flavescens L., (NW). Cyperus flavicomus Michx., (N). Cyperus gracilis R. Br., (NCI). Cyperus iria L., (TEN). Cyperus owanii Boeck, (NCI). Cyperus papyrus L., (NCI). Fimbristylis autumnalis (L.) Roem. & Schult., (NW). Scirpus cyperinus (L.) Kunth, (NW). Scirpus prolifer Rottb., (NCI). Eriocaulaceae Eriocaulon cinereum R. Br., (EXT). Hydrocharitaceae Limnobium laevigatum (Humb. & Bonpl. ex Willd.) Hei- ne, (NW). Iridaceae Tris foetidissima L., (N). Iris germanica L., (TEN). Tris orientalis L., (NCI). Ixia polystachya L., (NCI). Ixia speciosa Andrews, (NCI). Moraea collina Thunb., (N). Moraea polystachya Ker Gawl., (NCI). Juncaceae Juncus nodatus Cov., (N). Liliaceae (sensu lato) Agapanthus praecox Willd., (NCI). Allium cepa L., (NCI). Allium sativum L., (C). Amaryllis belladonna L., (TEN). Chlorophytum capense (L.) Druce, (C). HAyacinthus orientalis L., (C). Kniphofia uvaria (L.) Hooker, (NW). Leucojum aestivum L., (NCI). Narcissus pseudonarcissus L., (NW). Narcissus tazetta L., (NW). Ornithogalum umbellatum L., (GH/C). Pancratium maritimum L., (N). Tulipa clusiana DC. in Redoute, (TEN). Poaceae Acrachne racemosa (Roem. & Schult.) Ohwi, (TEN). Aira caryophyllea L. var. cupaniana (Guss.) Fiori, (NCI). 74 MADRONO Amphibromus neesii Steud., (NW). Aristida dichotoma Michx., (NW). Chloris truncata R. Br., (N). Echinochloa crusgalli subsp. spiralis (Vasing.) Tzvelev, (N). Echinochloa esculenta (A. Br.) H. Scholz, (NCI). Ehrharta longiflora Sm., (NW). Eragrostis curvula (Schrad.) Nees var. conferta Nees, (N). Eremochloa ciliaris (L.) Merr., (EXT). Gaudinia fragilis (L.) P. Beauv., (NW). Glyceria fluitans (L.) R. Br., (NW). Hordeum vulgare L. sensu lato, (C). Leptochloa dubia (Kunth) Nees, (N). Nassella tenuissima (Trin.) Barkworth, (C). Panicum maximum Jacq., (GH/C). Panicum repens L., (NCI). Panicum rigidulum Bosc ex Nees var. rigidulum, (NCI). Panicum texanum Buckl., (TEN). Pennisetum glaucum (L.) R. Br., (C). Pennisetum latifolium Spreng., (NCI). Phalaris coerulescens Desf., (C). Phyllostachys aurea A. & C. Riviere, (NCI). Phyllostachys bambusoides Siebold & Zuccarini, (NCI). Piptochaetium stipoides Hackel ex Arech. sensu lato, (NW). Polypogon imberbis (Phil.) Bjorkm., (NCI). Pseudosasa japonica (Sieb. & Zucc. ex Steud.) Makino ex Naka, (NCI). Schedonnardus paniculatus (Nutt.) Trel., (NCI). Spartina anglica C.E. Hubb., (NW). Stipa capensis Thunb., (NW). Themeda quadrivalvis (L.) Kuntze, (TEN). Tribolium obliterum (Hemzl.) Renvoize, (NW). Triticum aestivum L., (C). Pontederiaceae Heteranthera rotundifolia (Kunth) Griseb., (N). APPENDIX 2 ANNOTATED CATALOGUE Family circumscriptions and organization follow The Jepson Manual. Generic and specific applications reflect published treatments by specialists, modified only if clear- er information was gained by utilizing an alternative no- menclature. Abbreviations are as follows: DISTRIBU- TION (DIST) with geographic subdivisions as used in The Jepson Manual; CURRENT STATUS (CS); Current Status subcategories: NATURALIZED IN WILDLANDS (NW); NATURALIZED (OUTSIDE OF WILDLANDS) (N); PERSISTENCE TENUOUS (TEN); CASUAL (waif) (C); GREENHOUSE/CULTIVATION (GH/C); EXTIR- PATED (EXT); DOCUMENTATION (DOC). For details and descriptions of these categories see Tables 1 and 2. FERNS AND ALLIES Salviniaceae Salvinia molesta D.S. Mitch.: DIST: CCo, DSon, SCo: CS: NW, N (DSon), TEN (SCo): DOC: Imperial Co.: N side Hwy 98 in E Highline Canal nr. Winterhaven. T16S, R20E, Sec. 01, SB. Aug. 25, 1999, Johnson s.n. (CDA); All American Canal at Drop 1 nr. Coachella Canal. Sam- ple from equipment which at this point collects vegetation floating down the canal. Winterhaven area. T16S, R20E, Sec. 31, SB. Aug. 25, 1999, Johnson s.n. (CDA); River- side Co.: Drain canal of Palo Verde Irrig. District, Blythe [Vol. 49 region. TO6S, R23E, SB. Aug. 20, 1999, R. O'Connell s.n. (CDA); San Diego Co.: Private pond in vicinity of Fallbrook off Harris Truck trail. Oct. 28, 1999. S. Riviera s.n. (CDA); San Luis Obispo Co.: Private pond off Price Cyn. Rd. T30S, R12E, MD. Nov. 27, 2001, S. Stoltz s.n. (CDA, OBI); NOTES: Reported from San Diego River (San Diego Co.) Aug., 1999 but no specimen has been seen. Fallbrook location may have been purposely planted for increase and resale. Eradication attempts are currently underway by CDFA, USDA-APHIS in all sites. A Federal Noxious Weed, sale or growth is prohibited. Selaginellaceae Selaginella kraussiana (Kunze) A. Braun: DIST: NCo: CS: N: DOC: Humboldt Co.: Arcata. Damp, shad- ed areas near Humboldt State University Conservatory. April 5, 2000, G. Leppig 1286 (HSC); redwood forest be- hind Humboldt State University, along stream at Fern Lake. April 12, 2000, G. Leppig 1290 (CDA, HSC); Son- oma Co.: Best, C., et al. (1996, pg. 25). Ephedraceae Ephedra altissima Desf.: DIST: SCo: CS: NCI: DOC: San Diego Co.: Spreading from root shoots and climbing to 7 m to top of adjacent macadamia tree, site of old USDA Exp. Station, N of Science Park Dr., E of North Torrey Pines Rd, N of La Jolla. T15S, RO4W, Sec. O1, SB. Oct. 1, 1974, 7.C. Fuller 20049 (CDA). Ephedra distachya L.: DIST: SCo: CS: NW: DOC: Santa Barbara Co.: Smith, C.E (1998, pg. 372): NOTES: Reported as ‘established’ in Smith (1998), but by our criteria Wilken (SBBG) reports that it is naturalized in the oak woodlands about the Trout Club in the Santa Ynez Mtns. (pers. comm.). Det. by D. Wilken, verified also by S. Carlquist (SBBG). CONIFERS Pinaceae Pinus halepensis P. Mill.: DIST: CCo, SnFrB: CS: N: DOC: Contra Costa Co.: Mount Diablo, W slope Mount Zion, upper extent of Kaiser quarry reddish chert-like rocks on steep 25% W facing slope; mature trees (planted in rows), with abundant reproduction, juveniles to 10 ft tall. Elev. 1550 ft, TOIN, ROIW, Sec. 22, MD. Oct. 9, 1996, D.W. Taylor 15896 (UC); San Mateo Co.: Mc- Clintock, E., et al. (1990, pg. 62): NOTES: Seedlings are occasionally encountered about cultivated trees but are seldom allowed to mature. Pinus pinea L.: DIST: n Chl, SnFrB: CS: naturalized: DOC: Contra Costa Co.: Walnut Creek, Lakewood sub- division, on hillside. June 21, 1950, W.S. Malloch s.n. (UC); Santa Barbara Co.: Junak. S. et al. (1995, pg. 64). ANGIOSPERMS—DICOTS Acanthaceae Acanthus mollis L.: DIST: SnFrB: CS: C: DOC: San Mateo Co.: McClintock, E., et al. (1990, pg. 63): NOTES: Probably originating via garden waste; individ- uals sites often long-persistent, but permanence tenuous. Aceraceae Acer campestre L.: DIST: SnFrB: CS: N: DOC: Ala- meda Co.: lower Strawberry Canyon firetrail behind UC Berkeley campus, a few small trees at edge of woodland, 2002] June 11, 1998, B. Ertter & A. Rusev 16152 (UC): NOTES: Verbal reports indicate it may be sparingly nat- uralized elsewhere in the East Bay Hills. Acer saccharinum L.: DIST: ScV: CS: NW: DOC: Glenn Co.: Sacramento River at Butte City boat launch site, small tree on silty river bank. Oct. 22, 1998, Ertter 16448 (UC); Bank btwn. levee and W side of Butte Creek approx. % mi NE of confluence with Howard Slough, and immed. below McPherrin Dam (private and scheduled for removal). Two multi-trunked trees, + 12 meters tall, both apparently sterile, possibly persisting from old plantings. 59-2 TIN] P2153" We Sept 6s’ 1996hG.F* “Hrasa’ 13571 (CDA): Sacramento Co.: Sacramento River in Sacramen- to near the end of 10th St., elev. 85 ft. Apr. 28, 1997, D. Kelch DGK97.012 (UC); Betw. N side American River and William Pond, American River Parkway at Arden Way entrance. 38°33’N; 121°22’W. June 28, 1998, B. Mey- ers-Rice MR980601 (CDA, DAV); N bank American Riv- er, American River Parkway, few hundred meters down- river of the Estates Dr. entrance. June 28, 1998, B. Mey- ers-Rice MR980602 (DAV); American River Recreation Trad; E’ oi Union Pacific trestle. TO7N, ROSE, Sec. 31, MD. April 13, 1984, G.D. Barbe 4142 (CDA, DAV). Aizoaceae Galenia pubescens (Ecklon & Zeyher) Druce var. pu- bescens: DIST: SCo: CS: N: DOC: Los Angeles Co.: Ross, T. (1993, pg. 226-228); Ross, T. and S. Boyd (1996, pg. 432—433); Riverside Co.: U.S. Forest Fire Lab, Cyn. Crest Dr. El. 1200 ft. Fairly common in experimental planting of Artemisia californica, Eriogonum fasciculatum etc., Doubtless progeny of plants grown by E.C. Nord in 1970. Aug. 9, 1996, J. Beyers s.n. (UCR): NOTES: A specimen from San Diego Co. at UCR has not yet been accessioned. Amaranthaceae Amaranthus blitum L. subsp. emarginatus (Uline & Bray) Carretero: DIST: SCo: CS: C: DOC: Los Angeles Co.: waste ground at intersection of Beach Blvd. (Hwy 39) and Rosecrans Ave, at the Orange County line, La Mirada. Oct. 29, 1988, T. Yutani s.n. (CDA); Solitary small weed in a potted plant purchased at The Farm Store, Cal Poly Pomona. Jan. 10, 2002, A.C. Sanders 2489] (UCR): Riverside Co.: Moreno Valley, weed in untended planter beside Home Depot at Pigeon Pass Rd. and Hwy 160. Nov. 25, 2001, A.C. Sanders 24887 (UCR): NOTES: Determination by S. Mosyakin (KW, Mar. 2001), [=A. blitum subsp. polygonoides (Mogq.-Tandon) Carretero]. See Costea et al. (2001) for taxonomic discussion. Occa- sionally intercepted during nursery stock inspections on material from the US southeast, esp. Florida. Adapted to tropical and subtropical climates, this species would not be expected to be more than a minor or casual weed of greenhouse, garden or nursery. Sometimes treated (and reported) as A. lividus L. Amaranthus rudis Sauer: DIST: SCo, ScV: CS: N: DOC: Butte Co.: E side Sac. R. NW Parrott Landing, 1 mi. SE Ord Ferry, 12 mi. SW Chico. Riparian woodland. Sept. 21, 1999, L. Ahart 8267, 8266 (CDA, CHSC); 6 mi. W Chico, gravel bar, Sac. R. Sept. 26, 1981, L. Ahart 3205 (CAS, CDA, CHSC), det. by J.T. Howell 12/81; Sacra- mento Co.: NW corner Int. US 50 and Howe Ave., Sac- ramento. Single plant. Sept. 22, 1985, A. Shapiro s.n. (CDA, DAV); Santa Barbara Co.: SPRR, Carpenteria. Sept. 5, 1957, H.M. Pollard (CAS, CDA), det. J. Sauer 5- 1959 (as A. tamariscinus Nutt.); SPRR yards, Santa Bar- HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA > bara. Sept. 26, 1957, H.M. Pollard s.n. (CAS, CDA), det. J. Sauer 5/1959; loc. cit. Oct. 8, 1957, H.M. Pollard s.n. (DAV), det. J. Sauer 2/1974; SPRR yards, Santa Barbara. Oct. 8, 1957, H.M. Pollard s.n. (DAV): NOTES: Similar among dioecious California Amaranthi to A. arenicola. Pistillate plants readily distinguished among the California taxa by their two, rather than five pistillate sepals. Sta- minate plants of A. rudis have acute to acuminate sepal tips in contrast to the obtuse to retuse sepal tips in A. arenicola. Amaranthus tamariscinus Nutt. misapplied. Staminate and pistillate plants comprising Townsend s.n., Aug. 2, 1968, (CDA, RSA) collected nr. Pala in San Diego Co. and the source of the citation for A. tamariscinus in Munz (1974) USDA, NRCS (2001) and Beauchamp (1986), are, acc. to Hrusa, misidentified A. palmeri S. Wats. Brayulinea densa (Willd.) Small: DIST: SCo: CS: NCI: DOC: Santa Barbara Co: Howell, J.T. (1966, pg. 256); NOTES: Probably a roadside casual, not since re- ported in California. Froelichia gracilis (Hook.) Mog.: DIST: SCo: CS: NCI: DOC: Los Angeles Co.: Santa Fe RR, San Dimas. June 23, 1955, G.W. Garrettson s.n. (CDA). Det. by PA. Munz. Anacardiaceae Rhus lancea L.f.: DIST: SCo, DSon: CS: N: DOC: Riverside Co.: Coachella Valley, Palm Springs, W side of Palm Canyon Dr. just above (S of) junction with Hwy 111, weedy disturbed vacant lot, clearly spontaneous. 33°47.75'N, 116°32.52'W; TO4S, RO4E, Sec. 27, SB. EI. 140 m. Mar. 17 1996, A. C. Sanders and G. Helmkamp 17982 (UCR); Palm Springs, Murray Canyon, off Palm Canyon, a solitary arborescent shrub 4—5 m tall at the edge of the stream between palm groves, elev. 245 m, far from any cultivated plants, 33°45.5’N, 116°33’W, TO4N, RO3E, Sec. 10, SB. Feb 8, 1997, A. C. Sanders et al 19686 (UCR); Riverside, S side of the U.C.R. campus, a solitary shrub, apparently spontaneous, growing wedged between a large pecan tree and the wall of a concrete reservoir. Elev. 400 m. Feb 26, 1997, A. C. Sanders 19688 (UCR); Ventura Co.: Hills north of Moorpark, 0.5 mile west of Happy Camp Canyon, 1.9 miles north of Arroyo Simi Channel, TO2ZN R19W sec. 33, elev. 245-275 m, invasive in coastal sage scrub, particularly in small drainages. Jul. 27, 1995, C. Jones and R. Ramirez 8 (RSA, UCR): NOTES: Native to S. Africa and widely cultivated in Cal- ifornia, these are the first records of naturalized plants in California. Doubtless naturalized elsewhere in S. Calif. as the locations where it has been found represent much of the range of environmental conditions in lowland southern California, from the Sonoran Desert to the mild maritime influenced climate of Ventura County. Schinus polygamus (Cay.) Cabr.: DIST: SCo: CS: N: DOC: Los Angeles, Riverside, San Bernardino Cos.: Sanders, A.C. (1996, pg. 530): NOTES: Widespread in urban So. Calif., but is widely ignored by collectors. Still scarce in + natural areas, but often encountered in dis- turbed sites. Apocynaceae Nerium oleander L.: DIST: SCo, ScV: CS: NW: DOC: Los Angeles Co.: San Gabriel Mtns., in and near Glen- dora Wilderness Park, Harrow Canyon at the third debris basin. Elev. 1450 ft, Jul. 14, 1989, D. Swinney s.n. (UCR); Riverside Co: Temple, PJ. (1999, pg. 55); San Bernar- dino Co.: Waterman Canyon Road at old Arrowhead Hot 716 MADRONO Springs Resort, 7 km north Hwys 30 and 18 intersection. Sandy/rocky riparian with Artemisia californica and Vitis girdiana, May 9, 1972, E. Trubschenck 28 (UCR); Wa- terman Canyon, 1 km W of the old Arrowhead Hot Springs Hotel, 117°16'W, 34°11.5’'N, TOIN, RO4W, Sec. 11, SB. Elev. 1900 ft, abundant shrub in moist areas along the rocky canyon bottom along creek, numerous seedlings and plants of all sizes. Apr. 27, 1993, A. C. Sanders et al. 13824 (UCR); along Colorado River north of Parker Bridge, TOIN, R26E, Secs. 17 and 18, SB. Elev. 350 ft, mostly alkali and disturbed sites. May 2, 1978, Faulkner 572 (UCR).: Shasta Co.: Keeley, J., (1992, pg. 157): NOTES: Naturalized at the Waterman Canyon site for at least 30 years. Plants are abundant there, forming large thickets along the creek and obviously reproducing sex- ually. There is considerable variation in flower color, in- cluding shades of pink never seen in cultivation. Also es- tablished in nearby Hot Springs and Strawberry Canyons. The Swinney collection certainly represents a naturalized population as it shows (dried, no color notes) the mottled pink flower color that is common in the reproducing pop- ulation at Waterman Canyon. The Faulkner collection might be questioned because the dried flowers appear to be the pure white which is common in cultivated plants. There is nothing on the label which indicates that this collection was from cultivated or persisting plants, but there also is nothing eliminating that possibility. This re- port needs confirmation. Aquifoliaceae Ilex X attenuata Ashe: DIST: ScV: CS: NCI: DOC: Sacramento Co.: Volunteer tree on creek bank. Hoffman Lane, Fair Oaks. Jan. 30, 1977, Bly s.n. (CDA). Araliaceae Hedera canariensis Willd. (incl. H. algeriensis Hibb.): DIST: CCo, SCo, SnBR, SnFrB, SnGB: CS: NW: DOC: Alameda Co.: Albany Hill, abundant in oak forest. Jan. 14, 1995, B. Ertter 13918 (UC); Orange Co.: Trabuco Canyon. May 13, 1966, E.W. Lathrop 6297 (UCR); San Bernardino Co.: upper Waterman Canyon, San Bernar- dino Mtns. Dec. 23, 1998, A.C. Sanders 22369 (UCR); Dick Stoddard Canyon, San Gabriel Mtns. Jan. 15, 1994, Swinney 2695 (UCR); NOTES: Differs from H. helix in having larger leaves with fewer (most often only 3), more rounded lobes and fainter veins; more robust stems that are more shallowly rooted at the nodes; and rusty hairs on the growing tips that are basally fused into tightly ap- pressed, stellate-peltate trichomes (vs. white and + spreading in H. helix). Some naturalized forms not clearly separable from H. helix, needs further study. Occurring throughout the San Francisco Bay Area, where capable of being a serious pest plant; probably more common than H. helix in that region. Asclepiadaceae Asclepias fruticosa L.: DIST: deltaic GV: CS: C: DOC: Contra Costa Co.: Martinez Regional Shoreline, Granger’s Wharf Park, north end of Berrelessa St. Aug. 21, 1977, Walter and Irja Knight 3158 (CAS): NOTES: Narrow leaves like A. fascicularis, but pods ovate with long bristles. Not relocated in 2001, but site had been recently denuded by heavy grading. Calotropis procera (Ait.) Ait. f.: DIST: DSon: CS: C: DOC: Imperial Co.: agricultural area in the Imperial Val- ley. August, 1987, F. Laemmlen s.n. (UCR): NOTES: A [Vol. 49 solitary shrub, grower was concerned about its potential as a weed and is reported to have destroyed it after send- ing material for determination. Cynanchum louiseae Kartesz & Gandhi: DIST: SCo: CS: EXT: DOC: Riverside Co.: Sanders, A.C. (1996, pg. 526, 527); NOTES: The site of this population, which had been established for many years, was re-landscaped re- cently and the plants could not be relocated in 2001. Asteraceae Achillea filipendulina Lam.: DIST: NCo, SCo: CS: NCI: DOC: Humboldt Co.: dense growth in unkempt yard, McKinleyville. July 7, 1975, F. Bapeaux s.n. (CDA); Ventura Co: light infestation, 0.01 acre net over 1.0 acre gross, vacant lot, Poncho Rd., Camarillo. TOIN, R20W, Sec. 05, SB. June 20, 1979, H. Carpenter s.n. (CDA): NOTES: Gen. persistent and spreading vegeta- tively from cultivation, probably rarely reproducing by seed. Artemisia vulgaris L. sensu stricto: DIST: SCo, SNE: CS: NCI: DOC: Ventura Co.: Waste area of old habita- tion site between Ventura Ave. and Southern Pacific RR hear Wadstrom. Oct. 10, 1969, H.M. Pollard s.n. (CAS, CDA, SBBG); Mono Co.: Rock Creek Basin Rd. 0.1 mile N of Mono/Inyo Co. line, just S of Pine Grove Camp- ground, Inyo National Forest. TO5S, R30E, Sec. 31, MD. Aug. 12, 1981, G.D. Barbe 3532 (CDA): NOTES: De- terminations by S. Mosyakin (KW), 3/2001. Acc. to Mo- syakin, several other non-native taxa of the A. vulgaris alliance are represented at CDA but none can at present be associated with a specific name. Aster novae-angliae L.: DIST: KR: CS: NCI: DOC: Siskiyou Co.: Single plant in roadside ditch, escape from garden across the rd, Etna, Scotts Valley. Oct. 6, 1966. T.C. Fuller 15244 (CDA). Carthamus oxyacantha M. Bieb.: DIST: CCo: CS: C: DOC: Monterey Co.: Waif, screening disposal area, va- cant field south of spice processing plant, Schilling Place, Salinas. T15S, RO3E, Sec. 03, MD. Aug. 2, 1978. G.D. Barbe 2421 (CDA): NOTES: Carthamus oxyacantha is on the Federal Noxious Weed Act quarantine list (see Fed- eral Register, May 25, 2000, p. 33741-33743). Native to South Africa. Related species are aggressive invaders of pastures in New Zealand, Australia, California. Centaurea babylonica L.: DIST: n SNH: CS: C: DOC: Plumas Co.: Spontaneous at edge of lawn, to 6 ft tall, large rosette. County Hospital, Quincy. Aug. 7, 1972. F.H. Surber s.n. (CDA). Chrysanthemum balsamita L.: DIST: CaR: CS: C: DOC: Siskiyou Co.: volunteer, street side, Hennesy St., McCloud. Sept. 17, 1976, F.D. Horn s.n. (CDA). Cirsium scabrum (Poir.) Bonnett & Barratte: DIST: CCo: CS: NCI: DOC: Santa Cruz Co.: Howell, J.T, (1959, p. 27): NOTES: Collection made by A. Eastwood in 1900 and filed under the synonym Cnicus giganteus (Desf.) Willd. (UC). Acc. to J. Kartesz, this has also been reported as Cirsium giganteum (Desf.) Spreng. Coreopsis grandiflora Hogg. ex Sweet: DIST: CaR, SnFrB: CS: C: DOC: Alameda Co.: vacant lot, Appian Way, Union City. June 3, 1976, E. Whitaker s.n. (CDA); Plumas Co.: Disturbed roadside along Squirrel Ck, USFS road leading to Argentine Rock. Ca. 1 mi NE of Hwy 70, and ca. 7 mi E of Quincy. Elev. 1360 m. 39°55'N; 120°47'30"W. Aug. 22, 1996, G.F. Hrusa 13532 (CDA); San Bernardino Co.: a single waif, distinct from dwell- ings, San Bernardino Valley. June 10, 1909, S.B. Parish TA SI SIE): 2002] Cotula mexicana (DC.) Cabrera: DIST: CCo, ScV, SnFrB: CS: N: DOC: Alameda Co.: golf course green, Livermore. Apr. 20, 2000, E. de Villa s.n. (CDA); golf course green, Hayward. Apr. 26, 2000, G. Ingram s.n. (CDA); golf course green, Castro Valley. Apr. 26, 2000, G. Ingram s.n. (CDA); Marin Co.: golf course green, No- vato. Apr. 29, 2000, G. Ingram s.n. (CDA); Monterey Co.: golf course green, Fort Ord. Nov. 12, 2001, S. Fen- nimore s.n. (CDA, DAV); Sacramento Co.: golf course green, Elverta. Dec. 23, 1999. R. Chavez s.n. (CDA); golf course green, Sacramento. Jan. 21, 2000, V. Nyvall s.n. (CDA); golf course green, Sacramento. Jan. 9, 2000, F. Carl s.n. (CDA); golf course green, Galt. Feb. 9, 2000, D. Thompson s.n. (CDA); golf course green, Elk Grove. March 16, 2000, F. Carl, V. Nyvall s.n. (CDA); San Ma- teo Co.: golf course green, Pacifica. Mar. 21, 2002, D. Pendleton s.n. (CDA); Loc. cit., Apr. 18, 2002, (UCR); Siskiyou Co.: golf course green, Mt. Shasta region. Dec. 1, 2000, D. Smith s.n. (CDA): NOTES: A large infestation observed also in Napa Co. on a golf course near Pope Valley. Rapidly spreading perennial capable of competing and establishing by seed in mature low cut turf of greens and the adjacent collar. Not yet seen in taller mowed turf, or in wild situations, but expected in the latter. Easily overlooked due to diminutive stature. Native southern Mexico to Boliva, apparently at elevations above 3000 m. Crepis tectorum L.: DIST: SnBr, s SNH: CS: N: DOC: Inyo Co.: Mammoth Lakes, 37°38.8’N, 118°58.5'W, elev. 8100 ft. Uncommon in one local area on roadside among pines. Sept. 26, 1996, G. Helmkamp 1218 (UCR); San Bernardino Co.: Sanders, A.C. (1997b, pg. 307). Dendranthema X grandiflorum Kitam.: DIST: SCo: CS: NCI: DOC: Ventura Co.: heavy infestation in yard, Santa Paula. October 27, 1964, C.J. Barrett and V. Hol- mer s.n. (CDA): NOTES: This is the florist’s chrysanthe- mum. Original determination as Chrysanthemum morifol- ium Ramat. Dittrichia graveolens (L.) Greuter: DIST: CCo, SCo, ScV, SnFrB: CS: NW: DOC: Alameda Co.: San Francis- co Bay Wildlife Refuge at end of Cushing Road, forming dense masses on levee. Nov. 3, 1995, B. Ertter 14542 (UC); Shadow Cliffs Recreation Area, Livermore Valley, abundant in riparian woodland. Sept. 28, 1998, B. Ertter 16412 (UC); Merritt College, common in parking lot on west side of campus. Oct. 8, 2000, Ertter 17540 (UC): Gravelly sidewalk strip at corner of Monaco and Mission Drs. in Pleasanton. 37°39'08"N, 121°52’45”W. Nov. 22, 2000, D. Petersen 00-54. (CDA); Niles Cyn. Rd betw. Sunol and Fremont, mile marker 16. In Alameda Cr., sand/ eravel. bat,.37° 35.43" N,/121°54'22"W.. Oct: 27, 2001,-D. Petersen s.n. (CDA); Site M.5 storage area in Camp Parks, Dublin. Disturbed, graveled parking area. 37°43'28’N, 121°52'30"W. Oct. 17, 2001, D. Petersen 240 (CDA); Ar- royo de la Laguna at Verona Bridge (Pleasanton) on gravel bate” beside Stream. © 37757/35"N,, 121°52'55" W.- Nov:: 14, 2001, D. Petersen s.n. (CDA); Alameda/Contra Costa Co. [line]: Redwood Regional Park, West Ridge Road. Oct. 15, 2000, Ertter 17541 (UC): Contra Costa Co.: Lime Ridge, sterile flat in bowl of quarry area just south of Ygnacio Valley Road, in vicinity of Ygnacio Reservoir, Concord. Locally common. TOIN, ROIW, Sec. 08, MD. Oct. 18, 1998, B. Ertter and W.A. Morosco 16423 (JEPS): loc. cit. Dec. 29, 1999, Case and Ertter s.n. (CDA); San Mateo Co: About twenty plants on trailside, E end of Weeks St. adj. to Bay Lands Nature Preserve, E. Palo Alto. Oct. 18, 2001, J. Beall s.n. (CDA); Santa Clara Co.: Preston, R.E. (1997, 200-203); abundant, two miles north of Alviso Railroad tracks at upper edge of tidal HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA V7 marsh. Nov. 1, 1984, H.T. Harvey s.n. (CDA, SJSU, UC). Det. by C.W. Sharsmith, 5-88.; Overflow channel east of Coyote Creek, 25 yards W of Milpitas Sewage Treatment Plant. % acre. Oct. 16, 2000, J. Beall s.n. (CDA, UCR): roadside and in pasture at 4010 Calaveras Rd., Milpitas. Nov. 15, 2000, N. Garrison s.n. (CDA); Solano Co.: Sui- sun City, parking lot opposite wildlife center at Peytonia Slough, also near Civic Marina. Oct. 2, 2000, A.M. Sha- piro s.n. (DAV); Yolo Co.: City of West Sacramento, about 0.5 km W of Harbor Blvd., N of West Capitol Ave., along a jeep trail on the S side of the railroad embank- ment. Oct. 29, 1999, A.M. Shapiro s.n. (DAV): NOTES: Also noted by Ertter at Coyote Hills Recreation Area, and Lake Del Valle; by K. Hintsa at Rock City, Mount Diablo, all Alameda Co. D. Petersen (pers. comm. to Hrusa, 10- 2001), reports it is becoming common at Camp Parks, Alameda Co., where it occupies creekbanks as well as roadcuts and roadsides. A much-branched, densely glan- dular, odoriferous, fall-blooming annual, superficially reminiscent of a tarweed but with narrow overlapping phyllaries, or of Conyza but with yellow flowers. Ray flowers are reduced, and leaves are alternate. A rapidly spreading invasive weed. R. Preston (personal communi- cation to B. Ertter, 1-2000) reports it is now moving into the Central Valley as scattered individuals on most major highways leading inland from the San Francisco Bay Area. Emilia fosbergii D.H. Nicolson: DIST: SCo: CS: GH/ C: DOC: San Diego Co.: Weed in container grown nurs- ery stock. Pleasant Knoll Rd., Valley Center. Feb. 14, 2000, P. Nolan s.n. (CDA): NOTES: Has also been found as a rare nursery weed in Sacramento Co. (ScV) probably in imported soil. Potential garden weed. Emilia sonchifolia (L.) DC. ex Wight: DIST: SnJV: CS: GH/C: DOC: Tulare Co.: adventive under green- house bench; Terra Bella. T23S, R27E, Sec. 12, MD. May 28, 1987. R.D. Harris s.n. (CDA): NOTES: Potential gar- den weed. Grindelia papposa Nesom & Suh: DIST: SCo: CS: NCI: DOC: Ventura Co.: along RR in Ventura. 1962, H.M. Pollard s.n. (CAS, SBBG): NOTES: Reported in Smith, C.F, (1976, pg. 291) as Haplopappus ciliatus (Nutt.) DC., but excluded from the 2nd ed. (Smith, C.F, 1998) and perhaps not persisting. Helianthus tuberosus L.: DIST: CCo: CS: C: DOC: Alameda/Contra Costa Cos.: single individuals on both north and south banks of Cerrito Creek near end of Yo- semite Avenue at foot of Albany Hill. Sept. 30, 1995, B. Ertter 14526 (UC): NOTES: Source is presumably an up- stream creekside planting. Helichrysum petiolare Hilliard & B. L. Burtt.: DIST: CCo: CS: NW: DOC: Marin Co.: Matt Davis Trail above Stinson Beach, S side of Mt. Tamalpais, shade of forest, locally common in patches. July 25, 1992, B. Ertter and L. Fujii 11260 (JEPS); 200 m N of Panoramic Hwy, E of Stinson Beach, 200 m from the SW border of Mt. Tam- alpais State Park, growing near the remains of an old homestead, nr a larger population of plants 1 km distant. July 2, 1997, J. Randall s.n. (DAV); Monterey Co.: Open, sunny, sandy soil. Del Monte Forest on the edge of the 1959 burn, Monterey Peninsula. Growing through and up above manzanitas and other shrubs. Jan. 3, 1970, B. F. Howitt 3117 (CAS, CDA). Heliomeris multiflora Nutt. var. multiflora: DIST: SNE: CS: TEN: DOC: Mono Co.: Mammoth, vacant lot near the Post Office. Aug. 2, 1998, D.W. Taylor 16936 (UC). Determined by John Strother: NOTES: Apparently escaping from nearby areas seeded for ‘wildflowers’. The 78 MADRONO plants reseed in unmanaged, ruderal vegetation in the de- veloped portion of town, and in this setting reseed as an- nuals. Heliomeris multiflora var. nevadensis (Nelson) Ya- tes, a native perennial, occurs in the White Mountains to the east. Brought to our attention by D.W. Taylor. Helipterum roseum (Hook.) Benth.: DIST: SnFrB: CS: NCI: DOC: San Mateo Co.: McClintock, E., et al. (1990, pg. 79). Lasiospermum bipinnatum (Thunb.) Druce: DIST: SCo: CS: NW: DOC: Santa Barbara Co.: Ross, T. and S. Boyd (1996, 433-434). Leontodon muelleri (Schultz-Bip.) Fiori: DIST: ScV: CS: NCI: DOC: Glenn Co.: in alfalfa field, 6th and Wy- oming Aves, NE of Orland. T22N, RO2W, Sec. 08, MD. May 2, 1982, G. Stenlund s.n. (CDA). Oncosiphon piluliferum (L.f.) Kallersjo: DIST: SCo: CS: N: DOC: Riverside Co.: Sanders, A.C. (1996, pg. 528); Moreno Valley, E of Lake Perris along JFK Blvd. in disked field. March 23, 1998, R. Noll s.n. (OBI, SD). Det. by D. Keil, 11-2001; San Jacinto Wildlife Area, at headquarters off Davis Rd., north of Lakeview. Lakeview 7.5’ quad. 33°52'N, 117°07'W; T03S, RO2W Sec. 32, SB). Elev. 442 m/1450 ft. July 15, 2001, O.F. Clarke s.n. (UCR); San Jacinto Wildlife Area, Lovell Unit approxi- mately 885 m east of Davis Rd and 76 m north of the San Jacinto River levee. Lakeview 7.5’ quad, TO4S, RO2W, Sec. 05, SB. El. 433 m. Alkali playa with Rumex, Crypsis schoenoides, Phalaris minor, Atriplex argentea, etc. Trav- er loamy fine sand, saline alkali. June 18, 1995, D. Bram- let 2434 (UCR); San Jacinto Wildlife Area, approximately 920 m NW of Lakeview and 487 m west of the Davis Rd, 46 m N of Marvin Rd. Perris 7.5’ quad. TO4S, RO2W, Sec 06, SB. Elev. 433 m. Alkali playa with Plagiobothrys lep- tocladus, Crypsis schoenoides, Cressa truxillensis, etc. Willows silty clay. May 6, 1992, D. Bramlet 2265 (UCR); Romoland, on ramp to Hwy 215 from Hwy 74, just east of Hwy 215. Perris 7.5’ quad. 33°45’08"N, 117°11'06"W. Elev. 434 m/1425 ft. Uncommon on disturbed roadside, common in abandoned factory yard across 74 to the south. Also scattered along Hwy 215 all the way to Riverside. May 7, 2001, A.C. Sanders 24176, with Mitch Provance and T.B. Salvato (UCR); Val Verde, between Moreno Val- ley and Perris, along Hwy 215, 0.9 mi S of Oleander Ave., at S end of Patterson Ave. Steele Peak 7.5’ quad. 33°50'47"N, 117°15’05"W, TO04S, RO4W, Sec. 01, SB. El. 457 m/1500 ft. Weedy roadside on disturbed agricultural plains. Locally common on side of freeway, conspicuous for ca. 100 m. May 8, 2001, A.C. Sanders 24209, with Mitch Provance and T.B. Salvato (UCR); Moreno Valley, along I-215 just south of the March Field Museum, 0.7 mi S of Van Buren Blvd. Riverside East 7.5’ quad. 33°52'35"N, 117°15'52”W; TO3S, RO4W, Sec. 26, SB. EI. 465 m/1525 ft. Weedy roadside on disturbed plains. Lo- cally common and conspicuous for ca. 1 km. May 8, 2001, A.C. Sanders 24210, with Mitch Provance and T.B. Sal- vato (UCR); Lakeview Mtns., Pulsar View Rd, ca. 1 air- mile NE of Juniper Flats Rd. and ca. %4—1 mi by road from the base of the hills. Lakeview 7.5’ quad, 33°49'06’N, 117°05'W; T04S, RO2W, Sec. 15, SB. Elev. 610 m/2000 ft. Chaparral, burned within the past few years. Many na- tive wildflowers and some non-natives, all on NW-facing slope. Solitary ind. growing in open spot on the burn. Apr. 11, 1997, B. Pitzer 3121 (UCR): San Diego Co.: San Diego Wild Animal Park, off Hwy 78, E of Escondido in San Pasqual Valley. 33°06'N, 116°59’'W. Elev. 300 m/984 ft. Weedy area near back entrance to park. Few plants this year, previously common. Apr. 1998, Robert Noll, s.n. (UCR); San Diego Wild Animal Park, along back road [Vol. 49 behind exhibits (N of African Plains area); also seen on E edge of park and in S African section of Park Botanical Garden. 33°06’N, 116°59'W. Scattered to fairly common at edges of park; abundant among Aloes in S African sect. of Bot. Gdn. Elev. 300 m/984 ft. Mar 25, 1997, Jan Beyers s.n. (UCR): NOTES: Original report by Sanders was as the synonym Matricaria globifera (Thunb.) Fenzl in Harv. and Sond. Comment on the label of the Noll specimen above; “‘reported also from Orange Co., Haul Cyn. Rd., Irvine Ranch.”’ The current status of this population is not known. Osteospermum fruticosum (L.) Norl.: DIST: SCo: CS: NW: DOC: Los Angeles Co.: Zuma Beach area, mouth of Zuma Creek, E end of the County Beach. major veg- etation/habitats are small Salix lasiolepis stand w/under- story of escaped ornamentals; Typha/Scirpus marsh; rem- nant coastal dunes; disturbed roadsides, former parking area, and rubble dumping area. Site proposed for habitat enhancement. Occasional nr. stream. Point Dume 7.5’ quad. 34°01’N, 118°49'W. Elev. <25 ft. Mar. 12, 1997, S.D. White 4738 (UCR); Riverside Co.: Mockingbird Cyn. area, south of Van Buren Blvd. and ca. % air mi E of Mockingbird Cyn. Rd Riverside West 7.5’ quad. T03S, ROSW, Sec. 27, SB; 33°52'N, 117°23'W. Willow riparian with disturbed coastal sage scrub adjacent. May 8, 1989, Ed LaRue s.n. (UCR); NW Palomar Mountains, Agua Tib- ia Mountains; NW foothills of Dorland Mtn.: S end of Los Caballos Road, UC Emerson Oaks Reserve, at the Emerson Cottage. TO8S, RO2W, Sec. 24, SB. Elev. 1720 ft. Localized escape in Carpobrotus plantings from culti- vated plants around the Emerson Cabin. May 2, 1996, Darin L. Banks 0953, with E.H. Banks (UCR); San Luis Obispo Co.: Toro Canyon Rd., E of U.S. Hwy 101, just north of Cayucos. Disturbed soil in farmland. El. 400 ft. Aug. 6, 1989, G. Helmkamo s.n. (UCR); Santa Barbara Co.: Ferren, W. R., Jr. (1985, pg. 236): NOTES: The “freeway daisy”’ of commerce. Similar to O. ecklonis, but growth spreading rather than upright. Pericallis cruenta (DC. non Roth) Webb. & Berth.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 149); Thomas, H. (1961, p. 374); San Mateo Co.: McClintock, E., et al. (1990, pg. 83): NOTES: Cited reports are as Senecio cruentus DC. non Roth. Porophyllum ruderale (Jacq.) Cass.: DIST: SCo: CS: C: DOC: Orange Co.: Weedy areas along roads surround- ing vegetable crop field adjacent to Seal Beach Naval Weapons Station. Oct. 17, 2000. P. Guerrero s.n. (CDA): NOTES: First report for California of this widespread weedy plant, but probably casual as an escape from cul- tivation. Native (apparently) from Arizona, New Mexico, Texas to South America. Our material referable to the var. macrocephalum (DC.) Crong. (P. macrocephalum DC.). Apparently used medicinally and perhaps cultivated in southern California for that purpose. Ratibida columnifera (Nutt.) Wooton & Standl.: DIST: CaRH, SCoRI: CS: NCI: DOC: Lassen Co.: Growing in dry, rocky soil on the west side of Eagle Lake, two miles N of the Eagle Lake Resort on service road 33NO1. July 21, 1969, N. Santamaria 721 (CDA, CHSC); In re-growth vegetation in a cleared area at the north end of the Glenn-Eagle airstrip, southwest corner of Eagle Lake, 0.8 mi north of Eagle Lake Resort. T31N, RIOE, Sec. 04, MD. Aug. 20, 1974, G.D. Barbe 1939 (CDA, DAV); Riverside Co.: Murrieta, Warm Springs Creek, ca. ¥% mile south of Murrieta Hot Springs. 33°35’N, 117°08’'W, TO7S RO3W, SB. Elev. 1500 ft. June 8, 1995, G.R. Ball- mer and K.T. Stockwell s.n. (UCR), det. by A.C. Sanders, 2002] 2002.; June 5, 1996, K.T. Stockwell s.n (UCR); Jurupa Mountains: Glen Avon Heights, on the N side of Conning St. between Campbell and Lindsay Sts. Fontana 7.5’ quad. TO2S, RO6W, Sec. 03, SB. 34°01'35’N, 117°29'45’W, Elev. 860 ft Sandy alluvial slopes. Organic crust which was observed at this location two years ago is no longer present. Solitary perennial. July 30, 2000, M. Provance 2173 (UCR): San Benito Co.: San Benito. July, 1925, Mrs. J.A. Bettys s.n. (JEPS). Senecio squalidus L.: DIST: SnFrB: CS: NCI: DOC: Alameda Co.: volunteer plants abundant in an unculti- vated corner of the experimental growing grounds, U.C. Botanic Garden, Strawberry Canyon, Berkeley. Aug. 20, 1971, T.C. Fuller 19927 (CDA, DAV); north of corpora- tion/greenhouse area, UC Botanic Garden, Strawberry Canyon, Berkeley. May 17, 1979, G.D. Barbe 2511 (CDA). Tragopogon hybridus L.: DIST: CCo: CS: NCI: DOC: Monterey Co.: few plants as waifs, vacant field, Schilling Company, Salinas. T15S, RO3E, Sec. 3, MD. May 31, 1978, J. Lyons and B. Oliver s.n. (CDA). Tripleurospermum maritimum (L.) W. D. J. Koch: DIST: NCo: CS: TEN: DOC: Mendocino County: Man- chester, at the west end of town on gravel road to the beach, near north edge of Lagoon Creek T13N R17W, Sec 25, MD. Pt. Arena 7.5’ quad. El. 80 ft, 39°58'02’N; 123°42'08"W. Dominating a storage area for dairy silage. June 30, 2000, D. W. Taylor 17473 (JEPS): NOTES: =Matricaria maritima L. Brought to our attention by D.W. Taylor, det. by J. Strother (UC). Balsaminaceae Impatiens balfouri J.D. Hook.: DIST: n CCo, NCo, ne sCo, SakrB: CS: N: DOC: Wilken, D- (1993, pg. 2): NOTES: Included in supplement to Munz but accidentally deleted from Jepson Manual manuscript. Impatiens noli-tangere L.: DIST: n NCo, SnFrB: CS: N: DOC: Wilken, D. (1993, pg. 2) also: Alameda Co.: Strawberry Creek just above Haas recreation facility be- hind UC-Berkeley, local colony in moist shade, Aug. 29, 1996, B. Ertter 15271, (UC): NOTES: Included in sup- plement to Munz (as /. occidentalis) but accidentally de- leted from Jepson Manual manuscript. Berberidaceae Berberis darwinii Hook.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, pg. 73): Thomas, H. (1961, pg. 172): NOTES: Local escape from cultivation. Betulaceae Alnus cordata (Loisel.) Duby: DIST: NCoRO: CS: C: DOC: Sonoma Co.: Best, C., et al. (1996, pg. 87): NOTES: Establishing from landscape planting. Bignoniaceae Campsis radicans (L.) Bureau: DIST: deltaic GV: CS: TEN: DOC: Contra Costa Co: Antioch Marina, edge of marsh along roadside. July 11, 1998, B. Ertter and W.A. Morosco 16370 (UC); Sacramento Co.: brushy roadside along Hwy 160 approx. 1 mile S of Freeport and immed. N of Freeport Marina. Twining in Vitis californica, Rubus discolor. Not obviously persisting from cultivation. June 9, 2000, G.F. Hrusa 15440 (CDA, DAV): NOTES: A scattered escape from cultivation, possibly marginally nat- uralized? HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 79 Catalpa bignonioides Walter: DIST: n SNE SCo. ScV. SnJV: CS: NW: DOC: Amador Co.: Forster Ranch, along Dry Creek, just east of the Sacramento and San Joaquin Co. lines, 38°19’N; 121°00'W, elev. 50 ft, naturalized. June 6, 1990, L. LaPré s.n. (UCR), det. by A.C. Sanders: Fresno Co.: Kings River area, east of Centerville, just S of State Hwy 180 (Kings Cyn Rd.), along river, riparian forest and adjacent annual grassland, solitary tree ca. 4 m tall at forest margin near road, perhaps a persisting orna- mental planting, though there is no sign of a historic build- ing, 36°43'30"N; 119°28'00’W, T14S, R23E, Sec. 09, MD, elev. 350—400 ft. Apr. 3, 1997, S. White 4871 (UCR), det. by A.C. Sanders; Kern Co.: Bakersfield, E side of the canal at E end of Hart Memorial Park, in a dense thicket of willows, obviously spontaneous. April 25, 1959, E. Twisselman 5085 (DAV): Ventura Co.: Spontaneous in ditch on SPRR ROW at Loma Dr. crossing, Ojai Valley. May 29, 1971, H.M. Pollard s.n. (CAS, CDA): NOTES: Plants of uncertain status (probably persisting from culti- vation) are occasionally found in moist areas in western Riverside Co. (e.g., French Valley S of Winchester and San Jacinto River above Cranston Guard Station): Plants of uncertain status observed by Hrusa in Sacramento Co. along the American River at American River Parkway and in Yolo Co. along Sacramento River 8 mi S of Woodland. Expected elsewhere in low elevation riparian habitats. Boraginaceae Echium lusitanicum L.: DIST: CCo, NCo: CS: TEN: DOC: Monterey Co.: Rt 68, 1.5 miles south of Pacific Grove, occasional on roadside, May 22, 2000, G. Leppig 1380 (CDA, HSC): San Mateo Co.: common on Rt. 101 roadside and coastal scrub between Montara and Linda Mar. May 24, 2000, G. Leppig 1384 (CDA, HSC): NOTES: Also observed by Leppig as occasional on Rte. 1 in Mendocino Co. and on banks of the Gualala River in Gualala (Sonoma Co.). Pentaglottis sempervirens (L.) Tausch ex Bailey: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell. iE ctoak (1958; po: 11S): Thomas, Hoole ps 2388): NOTES: Reported as Anchusa sempervirens L. Brassicaceae Brassica fruticulosa Cyrillo: DIST: SCo, SnFrB: CS: N: DOC: Los Angeles, Riverside, San Bernardino Cos.: Sanders, A.C. (1996, pp. 523-524); San Mateo Co.: McClintock, E., et al. (1990, pg. 90). Cardamine flexuosa With.: DIST: CCo, SCo: CS: N: DOC: San Diego Co.: Vincent, M.A. (1997, pp. 305— 306): San Francisco Co.: In plantings nr. Fisherman’s Wharf. April 8, 1998, M.A. Vincent and E.H. Fried 8186 (DAV): NOTES: Most common as a greenhouse/nursery weed. Coincya monensis (L.) Greuter & Burdet: DIST: NCo: CS: NW: DOC: Humboldt Co.: Lot ESE of Manila Community Services District sewage pump station: dune sand mixed with gravel fill. Disturbed empty lot with Se- necio vulgaris, Lupinus arboreus, Holcus lanatus, Ra- phanus sativus, Eriogonum latifolium. Eley. 20’. 40°51'N; 124°10'W. Feb. 13, 1997, J. Belsher 2 (CDA, DAV, HSC): NOTES: Under eradication. Confirmed still present in 1999 (A. Pickart, personal communication). Iberis umbellata L.: DIST: CCo: CS: C: DOC: San Francisco Co.: Howell, J.T. et al. (1958, pg. 77). Thomas, H. (1961, pg. 185). Rorippa sylvestris (L.) Besser: DIST: SCo, SnJV: CS: TEN: DOC: San Diego Co.: Nursery property, Sidonia 80 MADRONO Rd., Encinitas Ca. Present for several years in this site. May 22, 1998, C. Elmore s.n. (CDA, DAY); loc. cit. June 28, 1998, J. Blasius s.n. (CDA): NOTES: All known sites under eradication. Also confirmed from commercial nurs- ery properties in San Joaquin Co. (Lodi), June, 1998; Ventura Co. (Oxnard), Jan., 2000. but submitted material not adequate for vouchering. Reproduction by root sprouts only but plants highly persistent; introduced and spread via contaminated nursery stock. Cabombaceae Cabomba caroliniana A. Gray: DIST: SnJV: CS: NW: DOC: Contra Costa Co.: Screen trap at Clifton Court Forebay, head of California Water Project. TOIS, RO4E, Sec. 35, MD. Oct. 18, 2000, R. Gage s.n. (CDA); San Joaquin Co.: Disappointment Slough NW of Stockton, TO2N, ROSE, Sec. 09, MD. Sept. 28, 1988; Griffin et al. s.n. (CDA); loc. cit. Oct. 6, 1995, Griffin, Finley s.n. (CDA); S edge 14 Mile Slough, abundant, with Egeria densa, Myriophyllum spicatum. TO2N, ROSE, Sec. 23, MD. Sept. 19, 1991, E. Finley, R. Villareal s.n. (CDA); Middle River about Bullfrog Marina. July 24, 2001, F. Maly s.n. (CDA): NOTES: Reported as common in Lew- iston Lake, Trinity Co. (DiTomaso personal communi- cation), but no confirming specimen or other plant mate- rial has been seen. Recognized by its submerged, deeply divided fan-shaped foliage leaves on distinct petioles. Emersed leaf-like bracts subtending inflorescences at the water surface are oval-perfoliate. Can fill at least a 3 meter water column. A purple-foliaged form is known and al- though not yet reported for California it is sold in the aquarium trade and is expected. Present in Disappointment Slough since at least 1980 [L. Anderson (USDA) personal communication]. Apparently spreading rapidly in the Sac- ramento Delta and ultimately to be expected widely. Vi- sual reports need verification as this species has been con- fused on cursory observation with Ranunculus aquatilis from which it differs in its perennial habit and petiolate fan-shaped leaves. Campanulaceae Campanula medium L.: DIST: SnBR: CS: C: DOC: San Bernardino Co.: San Bernardino Mtns, N side of Baldwin Lake, at Big Bear Landfill. Former pinyon-juni- per woodland, now disturbed and cleared; vegetation weedy except at lower edge, where it is Chrysothamnus nauseosus scrub. Solitary plant. Big Bear City 7.5’ quad., 34°18'40’N; 116°49'00"W; TO3N, RO2E, Sec. 30 and Sec. 31, SB. July 3, 2000, J. Wear s.n. (UCR), det. by A.C. Sanders. Lobelia erinus L.: DIST: SCo: CS: C: DOC: Marin Co.: Howell, J.T. (1970, p. 357); Santa Barbara Co.: Escape from cultivation on Cold Spring Road south of and near La Paz Road intersection, Montecito. June 16, 1965, H.M. Pollard s.n. (CDA, RSA, SBBG). Caprifoliaceae Leycesteria formosa Wallich.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 130-131); Thomas, H. (1961, p. 327): NOTES: Reported as a local escape from cultivation. Viburnum tinus L.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 131); Thomas, H. (1961, p. 327): NOTES: Local escape from cultivation. [Vol. 49 Caryophyllaceae Silene pseudatocion Desf.: DIST: CCo: CS: NCI: DOC: Monterey Co.: Howitt, B.E and J.T. Howell, (1973, pg. 14); road shoulder, old road of Hwy 1, Marina, Calif. March 13, 1970, A. Allison s.n. (CAS, CDA), det. by J.T. Howell, 6-4-1970. San Mateo Co.: Pacifica prop- erty, Redwood City. Mar. 9, 1961, A. Jillson s.n. (CDA), det. by N. van Kleeck, 3-1971; San Francisco Co.: Single plant, obviously spontaneous, in neglected weedy garden, San Francisco. April 19, 1972, J.T. Howell 48673 (CAS, CDA, DAY). Celastraceae Maytenus boaria Molina: DIST: SnFrB: CS: NW: DOC: Alameda Co.: Gwinn Canyon in Oakland Hills, north side of Marlborough Drive, regrowth of scrub after major fire. June 21, 1994, B. Ertter 12838a (UC); loc. cit. Aug. 29, 1997, E. Leong s.n. (UC): NOTES: A well- known ornamental small tree with drooping branches, nar- rowly rhombic, toothed leaves, and inconspicuous flowers. Considered a pest plant locally, with control efforts un- derway. Germinates profusely following fire. Chenopodiaceae Atriplex muelleri Benth.: DIST: DSon: CS: NCI: DOC: Riverside Co.: 20 miles west of Blythe, a single plant on roadside. Oct. 17, 1965, J.C. Roos s.n. (COLO, UC, UCR): NOTES: Perhaps extirpated, sporadic search- es over past 10+ years by Sanders have not found this. Chenopodium watsonii A. Nels.: DIST: SnGB: CS: NCI: DOC: Los Angeles Co.: Big Tujunga at Colby Ranch Rd. Elev. 3200 ft. Dry sunny sandy roadside. Oct. 1, 1966, L.C. Wheeler 8941 (CDA, RSA): NOTES: De- termination by S.E. Clemants (BKL) and S. Mosyakin (KW), 3-2001. Original determination as C. album L., dis- tinguished readily by the whitened reticulate seed coat of C. watsonii. Native in the Great Basin region, further col- lections and distribution data are necessary to determine if this is introduced or an overlooked California native. Salsola kali L. subsp. pontica (Pallas) Mosyakin: DIST: DMoj, s ChI: CS: NCI: DOC: Kern Co.: Mouth of Red Rock Cyn. Plant yellow-green, growing in shady probably sub-alkaline soil in dry wash. Occasional—com- mon in such places. Elev. 2300 ft. Creosote bush associ- ation. Aug. 22, 1961, E.C. Twisselmann 6468 (DAV). Det. by S. Mosyakin 3-2001. Ventura Co.: Mosyakin, S.L. (1996, pg. 389): NOTES: Ventura County record is a sin- gle collection from San Nicolas Island. U.S. Naval Radio- logical Defense Laboratory, near road above sand spit at 100 feet elevation, R. E. Foreman 42 (US). Label data provided by S. Mosyakin (KW). Cistaceae Halimium lasianthum (Lam.) Spach: DIST: SCo: CS: NCI: DOC: Ventura Co.: escaped in yards in Oxnard. June, 1949, V. Holmer s.n. (CDA): NOTES: Det. by M.K. Bellue. Convolvulaceae Calystegia silvatica (Kit.) Griseb. subsp. disjuncta Brummitt: DIST: NCo, SnFrB: CS: N: DOC: Alameda Co.: Codornices Creek at southwest corner of Albany Vil- lage. May 22, 2000, M. Hurlbert s.n. (JEPS); Humboldt Co.: Growing in waste area at the intersection of 11th and B Sts, Arcata. Moist gulch. Aug. 10, 1976, T. Nelson 3166 2002] (CDA); Arcata, 11th and B. St., empty lot. July 8, 2001, G. Leppig 1577 (CDA, HSC); Eureka, Waterfront Dr. waste area, 100 m N Humboldt County Library. July 10, 2001, G. Leppig 1578 (CDA, HSC); Arcata, South G St. adj. to city of Arcata Corporation Yd. Waste area, road- side. July 11, 2001, G. Leppig 1579 (HSC, UC); Marin Co.: Brummitt, R.K. (Madrono, in press): NOTES: Pre- viously confused with C. sepium subsp. limnophila (Greene) Brummitt, but readily distinguished by its larger flowers (5—7.5 cm), inflated bracteoles that hide the calyx, larger leaves, and glabrous vestiture. Sporadic but persis- tent in urban waste areas around Humboldt Bay. Convolvulus tricolor L.: DIST: CCo: CS: NCI: DOC: Monterey Co.: adventive, vacant field adjacent to Schil- ling Company processing plant, Salinas. T15S, RO3E, Sec. 03, MD. April 24, 1979, J.L. Johnson and B. Oliver s.n. (CDA), det. by G.D. Barbe, April 1979 at CAS. Dichondra micrantha Urb.: DIST: n SNE SCo, Sn- FrB: CS: NCI: DOC: Butte Co.: Heavy infestation, 1 acre, | mi E of Quincy Rd. and | mi S of Middle Fork Feather River on W side of Bidwell Mtn. Private property. June 7, 1963, W. Hansell s.n. (CDA), det. by T.C. Fuller; Los Angeles Co.: El Segundo Dunes, W of Los Angeles Int. Airport and Pershing Dr. 33°56'N, 118°26’W. Elev. 125 ft. Sept. 18, 1987, A.C. Sanders 7367 (UCR); Riv- erside Co.: Hemet, SE corner of State St. and Bibbel, 1712 ft hill at east end of Diamond Valley. Hemet 7.5’ quad. 33°42'N, 116°58'’W; TO5S, RO1W, Secs. 26, 27, 34, 35, SB, common corner. Elev. 494-518 m/1620—1700 ft. Scarce at margins of dried pool and in disturbed soil. May 3, 2001, A.C. Sanders 24113 (UCR); San Bernardino Co.: San Bernardino Mtns., E of Yucaipa, Water Cyn., trib. of Wildwood Cyn. from the N. Vicinity of old Hunt Ranch, ca. % mi. N of Wildwood Canyon Rd. (T02S ROIW Sec. 04, SB. 34°01'30"N, 116°59'30’W) Elev. 3300 ft. Fairly common on roadside at edge of barren cattle corral where clearly naturalized. June 5, 1992, A.C. Sand- ers 12365, with E.J. Lott and D. Pendleton (UCR); Ven- tura Co.: Flood plain of Coyote Creek nr. confluence with Ventura River, Foster Park. Oct. 15, 1969, H.M. Pollard s.n. (DAV): NOTES: Also noted on embankment of Cer- rito Cr., spreading beyond lawn of Creekside Park (Con- tra Costa Co.). Expected elsewhere. Cited in Munz (1974, pg. 379) as Dichondra repens Forst. & Forst.f. which is misapplied in California to D. micrantha, the common cultivated lawn substitute. Dichondra repens has also been misapplied in California to the native Dichondra donnelliana Tharp & Johnston, thus care must be taken, specifically a specimen observed, when applying an epi- thet to a reported occurrence identified as D. repens. Again, the importance of documenting voucher specimens is clear; in this case the holotype of Dichondra donnelli- ana was originally determined as D. repens! Ipomoea aquatica Forssk.: DIST: ScV: CS: C: DOC: Sutter Co: Weed in cultivated, diverse, vegetable row crop. Wet. Mung garden N of Yuba City. Oct. 1, 2001, G.F. Hrusa 15989 (CDA): NOTES: Becoming widely (and illegally) cultivated as a greenhouse crop in Califor- nia, less commonly as a row crop in the Central Valley, with weedy occurrences thus expected to increase in fre- quency. A potential aquatic pest in warm areas. Federal Noxious Weed. Ipomoea lacunosa L.: DIST: CCo: CS: C: DOC: San- ta Cruz Co: Aptos, watershed of Aptos Creek adjoining Nisene Marks State Park, weed along 3000 block of Red- wood Drive; elev. 550 ft, 37°00’45’”N; 121°54’00"W. Dis- turbed opening of residential yard in Sequoia sempervi- rens-Lithocarpus densiflorus-Quercus parvula vat. shrevei HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 81 dominated forest. Waif from birdseed originating from bird feeder. With waif Panicum sp. and Helianthus an- nuus. Sept. 6, 2000, D.W. Taylor 17585 (JEPS): NOTES: Perhaps trivial, but to be expected elsewhere. Originated from ‘National Aububon Society Superior Wild Bird Food’ [4.53 kg—Wagner Bros. Feed Corp.] purchased ca. spring, 2000. Native to eastern North America. The iden- tity of the Panicum sp. is unknown because it does not [has not] flowered. Ipomoea quamoclit L.: DIST: GV: CS: C: DOC: Sac- ramento Co: volunteer in residential garden, climbing on roses, Sacramento. April 17, 1992, Joe Bandi s.n. (CDA). Crassulaceae Crassula multicava Lem.: DIST: CCo: CS: NW: DOC: Monterey Co.: Pt. Lobos State Reserve. Rock crevices along the trail at Big Dome Cove, granite rock outcrops in Pinus radiata forest, with abundant Polypo- dium calirhiza and lichens, 36°32'01"N; 120°56'50’W, Feb. 2, 1998, D.W. Taylor 16282 (JEPS, UC), det. by Dean Kelch (UC); San Mateo Co.: McClintock, E., et al. (1990, pg. 102): NOTES: Reproducing by bulbils. This species was reported for Monterey County by Yadon (1995) but was not included in Matthews, M.A. (1997). Report and data provided by D.W. Taylor. Sedum album L.: DIST: c SNE SnFrB: CS: N: DOC: Tuolumne Co.: Yosemite Valley, Yosemite National Park, ruderal disturbed margin of paths and parking area in em- ployee housing tents on southeast edge of Yosemite Vil- lage area, disturbed margin of parking areas and paths in open sunny locations, 3900 ft, 37°44'35’N; 119°34'48"W. Sept. 9, 1997, D.W. Taylor 16266 (JEPS, UC): NOTES: Report and data provided by D.W. Taylor. Also observed spreading beyond cultivation into overflow basin in Creekside Park, El Cerrito (Contra Costa Co.). Sedum dendroideum Sesse & Moc. ex DC.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 80); Thomas, H. (1961, p. 188): NOTES: Local escape from cultivation. Cucurbitaceae Cucumis anguria L.: DIST: SnJV: CS: NCI: DOC: Madera Co.: in highway right-of-way, Madera. July 1939, M. Bellue s.n. (UC). Cucurbita ficifolia Bouche: DIST: SCo: CS: NCI: DOC: Ventura Co: Adventive on the waste ground of a once inhabited site betw. Ventura Ave. and So. Pac. RR nr. Wadstrom, Ventura Oil Fields. Nov. 27, 1968, H.M. Pollard s.n. (CDA). Cucurbita pepo L. var. medullosa Alef.: DIST: NCoRO: CS: C: DOC: Sonoma Co.: Best, C., et al. (1996, pg. 118): NOTES: The zucchini of commerce es- caping locally. Cuscutaceae Cuscuta reflexa Roxb.: DIST: SCo: CS: EXT: DOC: Los Angeles Co.: Abundant on Hedera canariensis, N side Biology Bldg., Cal State Los Angeles. October 23, 1969, 7.C. Fuller 19021 (CDA): NOTES: State listed Noxious Weed. Now eradicated, this was the only known occurrence in North America north of Mexico. Droseraceae Drosera aliciae Hamet: DIST: NCoRO: CS: NCI: DOC: Mendocino Co.: Meyers-Rice (Madrono, in press): restricted to its introduction site in a wet depression in a 82 MADRONO pygmy forest, 39°15'N; 123°45'W. Nov. 2, 1997, B. Mey- ers-Rice MR971101 (DAV): NOTES: Intentionally intro- duced. Drosera capensis L.: DIST: NCoRO: CS: NCI: DOC: Mendocino Co.: Meyers-Rice (Madrono, in press); sev- eral scattered colonies of naturalized plants spreading through wet depressions and seeps in a pygmy forest, 39°15'N; 123°45'W. Nov. 2, 1997, B. Meyers-Rice MR971103 (DAV): NOTES: Intentionally introduced. Ac- cording to B. Meyers-Rice (DAV) this species is probably D. linearis auct. non Goldie, as per Smith, G. and C. Wheeler (1990-1991, pg. 170) and Hickman, J.C. ed. (1993, pg. 541). Drosera tracyi MacFarlane: DIST: NCoRO: CS: NW: DOC: Mendocino Co.: Smith, G. and C. Wheeler (1990— 1991, pg. 170): NOTES: Intentionally introduced. Some- times treated within D. filiformis Raf. According to B. Meyers-Rice (DAV) both typical D. filiformis (as men- tioned in Hickman, ed. 1993) and D. tracyi are found at this site. Ebenaceae Diospyros virginiana L. var. virginiana: DIST: SnBr: CS: NCI Gn Mendocino Co.). NW (as clonal colonies in San Bernardino Co.): DOC: Mendocino Co.: Hopland Field Station, headquarters nr. office. June 14, 1959, AHM s.n. (AHUC); San Bernardino Co.: San Bernardino Mtns. Mill Creek Canyon, on S side of Hwy 38, 1.2 mi above Mountain Home Creek at Mountain Home Village, Forest Falls 7.5’ quad., 34°06'N; 116°58’30"W, TO1S, ROIW, Sec. 10, SB. Elev. 4040 ft/1232 m. Dry meadow w/scat- tered trees on alluvial bench in canyon bottom, sandy loam w/rocks. Single trees of Pyrus communis and Prunus cerasifera also present. Could be old orchard site, but veg- etation looked natural, w/no sign of former occupation; possibly a few spp. escaped from cultivation at Mountain Home. Grove of 74 trees, 2—9 m tall; oldest (75—100 yr) dead but 45 cm dbh, live trees 6-12 cm; corollas pale yellow, mostly w/4 lobes, but occasionally 5. Grove all female?, possibly from sprouts of old tree. Fruits pro- duced; but seeds? Discovered ca. 1983 by Goodman. Bark dark and deeply fissured into blocks, covered w/lichens. June 26, 1999, A.C. Sanders 22903 with John Goodman (ARIZ, CAS, MO, RSA, SD, UCR, UNLV, UTEP); loc. cit. Nov. 11, 1999, A.C. Sanders 23252 with Mihai Cos- tea, T. B. Salvato (UCR); Near Old Mormon Road Mon- ument on old loop off Hwy 18 below Crestline, San Ber- nardino North 7.5’ quad., TO2N, RO4W, Sec 27, SB. 34°13'30"N; 117°17'30"W. Elev. 4200 ft/1280 m, canyon woodland. Scarce tree ca. 20 m tall and 38 cm dbh. Road- side, presumably originally planted (possibly from dis- carded seed?), but with a number of saplings (10—15) de- rived from root-sprouts surrounding parent tree. Straight central trunk with small angled branches, square-checked bark. July 21, 2000, A.C. Sanders 23591 with N. Diep (UCR): NOTES: The Mendocino Co. specimen at AHUC may or may not document a spontaneous occurrence. Oth- erwise only two known populations but both are repro- ducing vegetatively and have been present for decades. Plants are vigorous and obviously successful under natural conditions. A population with both sexes present might be even more successful. Elaeocarpaceae Aristotelia chilensis (Molina) Stuntz [A. macqui LHer.]: DIST: SnFrB: CS: TEN: DOC: Alameda Co.: Strawberry Creek near Life Sciences complex on UC- [Vol. 49 Berkeley Campus, common shrub in understory along creek. May 23, 2000, B. Ertter s.n. (UC). Muntingia calabura L.: DIST: SnJV: CS: GH/C: DOC: Stanislaus Co.: Spontaneous in coco fiber imported from Sri Lanka, greenhouse hydroponic operation. Nov. 11, 1997, T. Watson s.n. (CDA): NOTES: Worldwide weed of the tropics and wet subtropics, indigenous to S. America. In California known only as a greenhouse weed (seedlings). May be expected to volunteer and persist un- der mild, moist conditions, esp. cultivated sites. Escalloniaceae Escallonia macrantha Hook. & Arn.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 81); Thomas, H. (1961, p. 192). Euphorbiaceae Euphorbia characias L.: DIST: CCo: CS: NCI: DOC: Alameda Co.: Albany waterfront, on former landfill, local colony (probably extirpated by subsequent park develop- ment). Aug. 19, 1994, B. Ertter 13076 (UC): NOTES: Single plant also noted on adjacent Albany Hill. Com- monly grown as an ornamental. Euphorbia cyathophora Murr.: DIST: SCo, SnJV: CS: NCI: DOC: Fresno Co.: to six feet, in roses, etc., Fresno. Oct. 28, 1959, T.C. Fuller 3213 (CDA); Ventura Co.: light infestation on roadside fill, Olsen Rd. betw. Thou- sand Oaks and Simi. Feb. 10, 1971, Hannah s.n. (CDA), det. by T-C. Fuller. Euphorbia dendroides L.: DIST: SCo: CS: NW: DOC: Los Angeles Co.: Sanders, A.C. (1997a, 203); Foothills, San Gabriel Mtns., Eaton Cyn nr. Kinneloa Mesa, Pasa- dena. TOIN, R12W, Sec. 13, SB. Mar. 30, 1981, S. Grang- er s.n. (CDA, RSA); Chantry Flats Ranger St., Big Santa Anita Cyn. Angeles Nat. Forest. May 21, 2002, J. Hart- man, s.n. (CDA, UC/JEPS, UCR); Santa Barbara Co.: Smith, C.E (1976, pg. 184); Santa Barbara, Franceschi Park, naturalized along rocky slope of Mission Ridge Rd. Sept. 5, 1950, C.F. Smith 2859 (DAV, SBBG); loc: cit Feb. 13, 1952, C.F. Smith 3224 (DAV, SBBG); Escaped on hillside, S of Franceschi Park, Santa Barbara. May 7, 1959, T.C. Fuller 2433. (CDA, DAV); Vacant field NE of Old Mission Santa Barbara. June 11, 1970, C.F. Smith 10229 (DAV, SBBG); Ventura Co.: Few scattered plants, Peto Seed Co. Ranch, Saticoy. May 20, 1959, 7.C. Fuller 2435 (CDA): NOTES: Several additional collections at DAV from the Franceschi Park locality, all from the 1950s, are not detailed here. Euphorbia heterophylla L.: DIST: ScV: CS: NCI: DOC: Sutter Co.: Weedy in 4 acre mung bean crop, pre- sent for past 2 seasons, 0.15 mi S of Nuestro Rd, W side Terra Buena Rd. NW Yuba City. Sept. 26, 1984, G.D. Barbe 4104 (CDA, DAV): NOTES: Reported also as an uncommon weed in the UCR (Riverside Co.) Botanic Garden. Euphorbia hirta L.: DIST: SCo: CS: N: DOC: Riv- erside Co.: Sanders, A.C. (1997a, 203-4): NOTES: Sometimes treated as Chamaesyce hirta (L.) Millsp. Has been intercepted as a weed of nursery stock from the southeastern states, esp. Florida. Euphorbia hypericifolia L.: DIST: CCo, ScV: CS: GH/ C: DOC: Monterey Co.: Town of Aromas, Blue Pacific Greenhouses at the corner of Carpenteria Rd. and San Juan Rd. Aug. 21, 2000, M. Inaba s.n. (DAV); Sacra- mento Co.: 12676 Stockton Blvd., uncommon ascending weed in greenhouse, presumably imported from Florida or Hawaii with foliage plants. May 23, 1988, D. Koutnik s.n. 2002] (DAV): NOTES: All original determinations as Chamae- syce hypericifolia (L.) Millsp. Euphorbia marginata Pursh: DIST: KR, n SNE ScV, SCo: CS: NCI: DOC: El Dorado Co.: S of Camino on rd to Pleasant Valley, adventive in roadfill. Aug. 7, 1977, G.L. Stebbins 77136 (DAV); Placer Co.: Limited light infestation, SPRR yards, Roseville. TION, RO6E, Sec. 11, MD. Aug. 26, 1971, Henderson s.n. (CDA); Shasta Co.: Limited heavy infestation, roadside, 100 ft N of Calif. Forestry Station, French Gulch, Clear Ck. Cyn. T33N, RO7W, Sec. 02, MD. Elev. 1500’. Sept. 8, 1970, P. Whipp s.n. (CDA); Ventura Co.: Spontanous in waste ground, Junipero St. betw. Santa Clara and Main Sts. Ventura. July 12, 1966, H.M. Pollard s.n. (CAS, CDA); spontaneous on ground cleared for waterfront development. Front and Palm Sts, Ventura. July 11, 20, 28, 1967, H.M. Pollard s.n. (CAS, CDA). Euphorbia myrsinites L.: DIST: TR: CS: NCI: DOC: Kern Co.: Single plant on stream bank, Vine Street, Fra- zier Park. TOON, R20W, Sec. 35, SB. May 14, 1981, J. Marks s.n. (CDA), det. by T.C. Fuller. Euphorbia rigida M. Bieb.: DIST: SnJV, SCo: CS: NCI: DOC: Tulare Co.: escaped from cultivation, RR ROW SE of Porterville. T22S, R28E, Sec. 06, MD. March, 1993, Ahrendes s.n. (CDA); Ventura Co.: 3—4 large plants established in grassy, weedy area, Erbs Rd., Thousand Oaks. March 30, 1967, Schall s.n. (CDA); in ice plant groundcover, median highway strip along US Hwy. 101 ca. 3 mi E of Ventura. Feb. 27, 1976, C. Elmore s.n. (DAV). Euphorbia terracina L.: DIST: SCo: CS: NW: DOC: Los Angeles Co.: Sanders, A.C. (1997a, 205): Volunteer in UCLA Botanic Garden. Oct. 23, 1967, T.C. Fuller 16495 (CDA); El Segundo Dunes, immed. W of LAX. May 18, 1988, A.C. Sanders 7832 (UCR, CDA); Solstice Canyon, Santa Monica National Recreation Area. TO1S, R18W, Sec. 16, SB. Mar. 22, 2001, S. Williams s.n. (CDA); Monterey Park. Garvey Reservoir, dry slope above dam. TO1S, R12W, Sec. 26, SB. June 26, 2001, J. Hartman and M. Adams s.n. (CDA); Palos Verdes Pen- insula, Rancho Palos Verdes, Ocean Trails development, between Palos Verdes Dr. South and the ocean. San Pedro 7.5’ quad. 33°43'37"N, 118°20'30’W. Elev. 2—10 ft. Base of coastal bluffs. July 17, 2001, Jeremiah George s.n. (UCR), det. by A.C. Sanders; Zuma Beach area, mouth of Zuma Creek, E end of the County Beach, ca. 100—200 plants on W bank of the stream, in sand, and on adjacent remnant dunes. Point Dume 7.5’ quad. 34°O1'N, 118°49’W. Elev. <25 ft. Mar. 12, 1997, S.D. White 4750 (UCR), det. A.C. Sanders: Santa Monica Mtns., Malibu Lagoon, Malibu Beach State Park, mouth of Malibu Creek. Malibu Beach 7.5’ quad. 34°02’N, 118°41'W; TO1S, R17W Sec. 32, SB. Elev. 8 m/25 ft. Fairly common perennial at edges of cultivated areas at Adamson house. Oct. 10, 1998, A.C. Sanders 22259 (UCR); same as pre- vious, but: uncommon at edge of road (Cross Creek) on W side of lagoon. Oct. 10, 1998, A.C. Sanders 22260 (UCR): NOTES: Monterey Park form may be at least facultatively annual, warrants further study. Sapium sebiferum (L.) Roxb.: DIST: ScV: CS: NW: DOC: Sacramento Co.: On the N bank of American Riv- er in the American River Parkway, few hundred meters downstream of the Estates Drive entrance. 38°33'N; 121°22’W. June 28, 1998, B. Meyers-Rice MR980603 (CDA, DAV): NOTES: Ornamental, commonly cultivated in the residential areas surrounding the American River Parkway. Has the potential to naturalize locally in Cali- fornia. Reported from SnFrB, but no confirming speci- HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 83 mens have been seen. A serious pest in the summer wet southeastern U.S. Fabaceae Astragalus cicer L.: DIST: c SNF: CS: TEN: DOC: Tuolumne Co.: Disturbed waste area nr. Standard. July 4, 1998, M. Chambers s.n. (CDA): NOTES: Persisting in this site for several years previous to 1998 and observed inel999.- Cassia nemophila A. Cunn.: DIST: DSon: CS: TEN: DOC: Riverside Co.: Coachella Valley, between La Quinta and Indio, along Jefferson St. between Ave 54 and I-10, 33°41'N; 116°16’W, TO6S, RO7E, Sec. 04, elev. 10 m. Fairly common 1—2 m shrub, scattered on roadside and in adjacent old fields, cultivated nearby in center divider of Jefferson St. and spreading by seed into adjacent dry lands. May 28, 1999, A.C. Sanders 22794 (UCR, and to be distributed): NOTES: Reproducing without care in one of the driest parts of the state. [This belongs in Senna, but as of 1986 the appropriate combination had not been pub- lished (R. Barneby, personal communication; Royal Hort. Dict. Gardening, 1992, says the same, but uses the Senna nemophila combination anyway “awaiting publication’’)]. Ceratonia siliqua L.: DIST: SCo: CS: NW: DOC: Los Angeles, Riverside, San Bernardino Cos.: Sanders, A.C. (1996, pg. 526). Coronilla valentina L.: DIST: SCo, s ChI: CS: NW: DOC: Los Angeles Co.: Ross, T. and S. Boyd (1996, pg. 435). Dolichos lignosus Pers.: DIST: SCo: CS: NCI: DOC: San Diego Co.: Beauchamp, R. M. (1986, pg. 157). Genista monosperma (L.) Lam, non Link, nec Del.: DIST: SCo: CS: NW: DOC: Los Angeles Co.: San Ga- briel Mtns, base of range at Padua Hills, W of mouth San Antonio Cyn. Disturbed alluvial fan and adjacent slopes with chaparral. Feb. 3, 1990, S. Boyd 3828 (CDA, RSA): San Diego Co.: South of Fallbrook. 100 yds W of Olive Hill Rd, approx. %4 mi S of Mission Rd. (rd $13), S side of Color Spot nursery. June 19, 2000, J. Giessow s.n. (CDA and to be distributed): NOTES: Original det. by E. McClintock (CAS); label det. of Boyd 3828 as Genista aetnensis (Biv.) DC. Treated in Flora Europaea as Lygos monosperma (L.) Heywood and listed by CalEPPC as Re- tama monosperma (L.) Boiss. We find the generic distinc- tions dubious. Under eradication on adjacent Camp Pen- dleton federal lands (personal communication from E. Johnson, May 2000). Specimen cited in Rejmanek and Randall (1994) as at DAV apparently was never deposited. Gleditsia triacanthos L.: DIST: GV: CS: NW: DOC: Sacramento Co.: Randall, J.M. and B. Meyers-Rice (1997, pp. 399-400). Lathyrus sativus L.: DIST: CCo, NCoRO: CS: C: DOC: San Luis Obispo Co.: Spontaneous in garbanzo bean field, opposite Los Osos Valley Memorial Park Cem- etery, W end of Los Oso Valley. July 22, 1971, J.H. Foott s.n. (CDA, DAV); Sonoma Co.: Best, C., et al. (1996, pg. ta): Ononis alopecuroides L.: DIST: SCoRO: CS: NW: DOC: San Luis Obispo Co.: Hrusa, G.F (2000, pg. 139): NOTES: Known from a single large population, currently under eradication by San Luis Obispo Co. Agricultural Commissioner’s Office. Robinia hispida L.: DIST: ScV: CS: N: DOC: Sac- ramento Co.: Colony on bank of Sacramento River at divergence of Steamboat Slough, SW corner of Steamboat Bridge. Root-sprouting, spreading from planted plants down bank, forming a thicket beneath Robinia pseudo- 84 MADRONO acacia. Apr. 9, 2000, G.F. Hrusa 15318 (CDA and to be distributed): NOTES: Locally naturalized, but apparently spreading only vegetatively. Robinia hispida consists of a series of clones, reproducing facultatively by rootsprouts and agamospermic seeds (Isely, 1998). Several forms are cultivated. Senna artemisioides (Gaudich. ex DC.) Randell: DIST: SnBr, SnGb: CS: N: DOC: Los Angeles Co.: San Gabriel Mts, north of Claremont, along Burbank Fire Road in Burbank Canyon, west of Palmer Canyon; grow- ing near edge of gravel road, naturalized in area. Jan. 24, 1993, 7.S. Elias 12445 (UC); San Bernardino Co.: North of San Bernardino along Hwy 18, 0.2 miles above the lower end of old Waterman Canyon Road, SW side of highway on road fill, growing wild. Mar. 9, 1984, F.C. Vasek s.n. (UCR): NOTES: Population still extant ca. 1995-1998 in lower Waterman Canyon just above Hwy 18. All specimens labeled as Cassia artemisioides Gau- dich. ex DC. Senna obtusifolia (L.) H.S. Irwin & Barneby: DIST: DSon, SnJV: CS: N in DSon; TEN in SnJV: DOC: Riv- erside Co.: Sanders, A.C. (1996, pg. 531): NOTES: A single individual also found on a roadside in Fresno Co., sent without additional data to CDA for confirmation; ma- terial in condition too poor for vouchering. Sesbania punicea (Cav.) Benth.: DIST: CaRE GV, NCoRO: CS: N: DOC: Butte Co.: Shrub on wet sand, margin of small pond, % mile W of Pacific Heights Rd., Oroville Wildlife Area. T18N, RO3E, Sec. 03, MD, elev. 140 ft. Aug. 23, 2000, L. Ahart 8660 (CDA, CHSC). Fres- no Co.: gravel pit ponds, Pinedale. June, 1988, J. Dun- nicliff s.n. (CDA); Sacramento Co.: On the margins of William Pond in the American River Parkway, Arden Rd. entrance. 38°33’N; 121°22’W. June 28, 1998, B. Meyers- Rice MR980604 (CDA, DAV); Shasta Co.: E side Hwy 273 in Redding, immed. S of Breslauer Rd. Shrubs in wash bet. Hwy and RR tracks. Aug. 21, 2000, K. Martyn s.n. (CDA); Riverview Country Club, Bechelli Lane. Ex- tensive lakeshore infestation. Dec. 11, 2001. K. Martyn s.n. (CDA); Sonoma Co.: Best, C., et al. (1996, pg. 140): NOTES: Sonoma County report as S. tripletii Host. Trifolium alexandrinum L.: DIST: CCo, ScV: CS: NCI: DOC: Butte Co.: South side of Evans-Reimer Rd., ca. 1 mi E of Pennington Rd., Gray Lodge Waterfowl Management Area. May 10, 2001, L. Ahart 8738 (CDA, CHSC, UC); Monterey Co: few waifs, vacant field. Schil- ling Co., Salinas. T15S, RO3E, Sec. 03, MD. May 31, 1978, J. Lyons and B. Oliver s.n. (CDA). Trifolium cernuum Brot.: DIST: ScV: CS: N: DOC: Butte Co.: Near boat ramp off Larkin Rd., Thermalito Afterbay, Lake Oroville. May 13, 2000, L. Ahart 8343 (CHSC, MU, UC/JEPS): NOTES: Determination con- firmed by M. Vincent (MU). Reported in Oswald, V. (2000). Naturalization local. Trifolium gemellum Poir. ex Willd.: DIST: CCo, SnFrB: CS: N: DOC: Napa Co.: Henry Road 1.7 miles northwest of Dealy Lane [SW of city of Napa]. Common on open, grassy, southwest-facing slope in Quercus agri- folia woodland, elev. 450 ft, TOSN, ROSW, Sec 12, MD. May 12, 2000, J. Ruygt 4248 (UC): NOTES: Naturali- zation local. Report and data provided by J. Ruygt. Trifolium resupinatum L.: DIST: NCo, CCo, ScvV, SCo: CS: NCI: DOC: Humboldt Co.: immediate vicinity of Eureka, a single plant. May 14, 1896 and July 8, 1897, J.P. Tracy 105 (UC); Monterey Co.: few scattered plants in vacant field, flowers pinkish; Schilling company, Sali- nas. T15S, RO3E, Sec. 03, MD. May 11, 1978, J. Bunch and B. Oliver s.n. (CDA); Santa Barbara Co.: Howell, [Vol. 49 J.T. (1972, pg. 103); Smith, C.E (1976, pg. 179); Sutter Co.: Edge of field, 3 miles S of Oswald. June 11, 1967, J.T. Howell 42556 and G.H. True (CAS, CDA); Ventura Co.: Howell, J.T. (1972, pg. 103): NOTES: Probably only casual. Trifolium retusum L.: DIST: CaRF: CS: NW: DOC: Tehama Co: Calif. Dept. of Fish and Game parcel on west side of Manton Road ca. 2 miles north of Dales Sta- tion on Hwy 36, ca. 14 miles northeast of Red Bluff, T29N, RO2W, Sec 26, MD. Elev. 740 ft, locally abundant in gravelly soil. May 20, 1996, V.H. Oswald and L. Ahart 7613 (JEPS); loc. cit. May 21, 1998, V.H. Oswald 9087 (JEPS): NOTES: Determination by M. Vincent (MU). Trifolium stellatum L.: DIST: CCo: CS: NCI: DOC: Monterey Co.: waif, vacant field, Schilling Company, Sa- linas. T15S, RO3E, Sec. 03, MD. May 31, 1978, J. Lyens and B. Oliver s.n. (CDA). Trifolium striatum L.: DIST: n SN: CS: NCI: DOC: Nevada Co.: Grass Valley, East Main Street at Dorsey Drive, waste ground at entrance to Litten Industries com- plex, TI6N, RO8E, Sec. 23, MD. June 3, 1980, G.D. Barbe and P. Hiatt 2791 (CDA, JEPS); Spring Hill be- tween Grass Valley and Nevada City. Elev. 2700 ft. June 19, 1973, G.H. True 7584 (CAS, CDA); loc. cit. May 11, 1973, G.H. True 7469 and J.T. Howell (CAS, CDA); Son- oma Co.: Best, C., et al. (1996, pg. 144); Sugarloaf Ridge State Park, ca. 1 air mi S of Red Mtn, Adobe Cyn. TO7N, RO6OW, Sec. 22, MD. May 21, 1996, F. Bowcutt 2141 (DAV). Trifolium tomentosum Willk. ex Nyman: DIST: NCoRO, ScV, SnFrB: CS: NW: DOC: Contra Costa Co.: Mount Diablo State Park, connector between Barbecue Terrace Road and Wall Point Road, localized patch at edge of path through open oak woodland/grassland. May 3, 1998, B. Ertter 16063 (JEPS); Napa Co.: Imola Ave. 0.2 mile west of Suscol Ave., Napa. Grasslands, elev. 20 ft, TOSN, RO4W, Sec. 14, MD. April 11, 1979, J. Ruygt 501 (JEPS); Sacramento Co.: Pasture adj. to underpass at Rio Linda and SW Elverta, betw. Natomas E Main drain and W end of ‘N’ St. TION, ROSE, Sec. 19, MD. Elev. 40 ft. May 12, 1992, R. York 92-002 (CDA); Overflow parking area, E of Cal Expo at Ethan Way, Sacramento. June 7, 1987, N. Wymer s.n. (CDA, DAV); Sonoma Co.: Best, C., et al. (1996, pg. 144); Annadel State Park, east of Ledson Marsh, several dense patches in grassland, both sides of Marsh Trail, midway between intersection with Lawndale Trail and marsh spillway. April 19, 2000, A. Howald 2037 (CDA): NOTES: Resembling T. fragiferum in having the calyx inflated in fruit, but annual and with only a vestigial involucre. All specimens at CDA appear referable to the var. tomentosum sensu Zohary and Heller (1984). Easily overlooked and probably more widespread than the collections above indicate. Trifolium vesiculosum Savi: DIST: CCo, NCoRO, ScV: CS: N: DOC: Humboldt Co.: Disturbed road repair area at roadside, Hwy 101 nr Orick. TOON, ROIE, Sec. 06, H. Sept. 14, 1998, P. Haggard s.n. (CDA); Santa Cruz Co.: Common on limestone mine tailings above Davenport. Nov. 26, 2000, G.F. Hrusa 15725 (CDA); So- lano Co.: SW of Davis, abandoned fields between Hwy 113 and Pedrick Rd. July 1, 1998, M. Rejmanek s.n. (CDA, DAV): NOTES: A distinctive species among na- tive and naturalized California clovers, readily distin- guished by its chartaceous, inflated calyces having 25+ prominent longitudinal veins and similarly prominent lat- eral venation. Apparently an occasional component of clo- ver-containing hydro-seed mixtures; Humboldt Co. and 2002] Santa Cruz Co. occurrences may have originated via this pathway. Trigonella corniculata L.: DIST: NCoR: CS: NCI: DOC: Mendocino Co.: Ukiah, in cover crop. May 5, 1938, G.T. Nordstrom s.n. (UC). Trigonella foenum-graecum L.: DIST: CCo, ScV: CS: NCI: DOC: Monterey Co.: Volunteer in vacant field next to Schilling plant, Salinas. T15S, RO4E, Sec. 03, MD. April 24, 1979, B. Oliver s.n. (CDA); Yolo Co.: north edge of Davis just west of B St., weed in barley field. Mar. 27, 1951, J.M. Tucker 2058 (DAV, UC). Vicia bithynica (L.) L.: DIST: CCo: CS: NCI: DOC: Monterey Co.: waste area, Schilling Co., Salinas. T15S, RO4E, Sec. 03, MD. May 15, 1978, B. Oliver s.n. (CDA). Fagaceae Quercus ilex L.: DIST: SCo: CS: N: DOC: Los An- geles Co.: Claremont, adventive in plantings of chaparral shrubs on grounds of Rancho Santa Ana Botanic Garden, 15 Nov. 1990, S. Boyd and T. Ross 5305 (RSA, UCR); Orange Co.: City of Orange, N end of Yorba St. at San- tiago Creek, near Chapman Ave. crossing. Orange 7.5’ quad. 33°47'24’N; 117°50'24’W, elev. 260 ft/79 m. Creek- bed, several trees to 30 ft. Feb. 1, 2000, Y. Moore s.n. (UCR): Riverside Co.: Weed tree in a citrus orchard in Rubidoux, area being cleared for houses, but this tree be- ing retained, Oct. 23, 1994, Donald E. Peck s.n. (UCR); Riverside, east side of UCR campus nr. parking lot 13, assoc. with Salix lasiolepis, Populus fremontii, Baccharis salicifolia, etc. Elev. 335 m/1100 ft. Solitary 5 m tree at edge of wash and parking lot. Clearly spontaneous—no cultivated plants in immediate vicinity. May 12, 1997, A.C. Sanders 20711 (UCR); Riverside, Watkins Dr. east of Blaine St., at Lemona Siding. Riverside East 7.5’ quad. 33°58'30"N, 117°19'W; TO2S RO4W Sec. 20, SB. Elev. 335 m/1100 ft. In hedge of oleander and Brachychiton along N side of Watkins. At least 6 young trees, mostly 3—5 m tall, growing as weeds in hedge; plainly sponta- neous. May 3, 2001, A.C. Sanders 24123 (UCR). Geraniaceae Geranium columbinum L.: DIST: NCo, ScV: CS: NCI: DOC: Humboldt Co.: ca 7.5 road miles south of Ferndale, along Wildcat Road near Green Pond ranch. May 20, 1987, Pykala and Norris 751 (MO); Solano Co.: 5.6 miles W of Winters. April 16, 1968, Ishizuka 19 (MA): NOTES: Acc. to Aedo (2000) this is a relative of G. car- olinianum L., native to the Old World but widely distrib- uted in the northeast US; also in Oregon and Washington. Geranium lucidum L.: DIST: SnFrB: CS: N: DOC: Alameda Co.: lower end Strawberry Canyon firetrail be- hind UC-Berkeley Botanical Garden, locally abundant in wet ground at edge of forest. Apr. 11, 1998, B. Ertter 15979 (UC); loc. cit. Apr. 29, 1998, Ertter 16029 (UC). Geranium purpureum Vill.: DIST: NCoR, SnFrB: CS: NW: DOC: Alameda Co.: lower end of Strawberry Can- yon firetrail, behind UC-Berkeley campus. Nov. 23, 1991, B. Ertter and B. Olson 10891 (UC); loc. cit. Apr. 25, 1974, L.R. Heckard 3679 (JEPS); Albany Hill, uncommon at time of collection but rapidly becoming more abundant. May 20, 1995, B. Ertter 14216 (UC); corner of Hearst Ave. and Gayley Road, UC-Berkeley campus. Apr. 4, 2001, B. Ertter and D. Norris 17574 (UC and to be dis- tributed); Napa Co.: Kroeber Ranch west of Rutherford, in weed-filled meadow. Apr. 25, 1996, B. Ertter and J. Ruygt 14601 (UC): NOTES: Similar to G. robertianum, but witn consistently smaller (<< 1 cm long), more uni- HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 85 formly bright pink petals, yellow (vs. orange) anthers, and less anthocyanic foliage overall. Geranium pyrenaicum Burm. f.: DIST: CCo: CS: C: DOC: Alameda Co.: Berkeley Campus, east side of Bot- any building. June 15, 1914, W.L. Jepson s.n. (JEPS): NOTES: Current determination by Ertter, 1996; previous- ly determined as (and possibly basis for CCo record of) G. pusillum Burm.f. in Hickman (1993). Geranium rotundifolium L.: DIST: CCo, SCo, SnFrB: CS: NW: DOC: Alameda Co.: UC-Berkeley campus, in Grinnell Natural Area, locally abundant. June 28, 1996, B. Ertter 14910 (UC); Albany Hill. July 13, 1996, B. Ert- ter 14967 (UC); Contra Costa Co.: Tilden Regional Park east of Grizzly Peak Boulevard, scattered colonies along dirt road in weedy hillside. May 14, 1996, B. Ertter 14684 (UC); Los Angeles Co.: Ross, T. and S. Boyd (1996, 435— 436); San Luis Obispo Co.: Huasna Rd., 1.5 rd miles E of bridge over Huasna River; foothill woodland and chap- arral patches, area grazed. Occasional on roadside. April 1, 1995, D. Keil 24714 (OBI); US 101 1.3 miles NW of Cuesta Pass, ca. 0.4 miles S of Tassajara Creek Rd.. Coast- al live oak woodland and adjacent roadside zone, locally common in roadside zone and edge of woodland under- story. May 27, 1999, D. Keil 28354 (OBI); Cuesta Grade East along Mt. Lowe Rd., ca. 2 road miles from Highway 101, locally common under shade of Quercus agrifolia on bank above road. May 9, 2000, D. Keil 28648 (OBI): NOTES: Somewhat reminiscent of Geranium molle, but with entire (vs. apically notched) petals, a short awn (< 1 mm long) on the sepals, fruits that are finely hairy rather than wrinkled, and less deeply lobed leaves. Noted as weed elsewhere in Berkeley; possibly seen by Ertter on Fremont Peak, San Benito Co. Geranium texanum (Trel.) A. Heller: DIST: CCo: CS: NCI: DOC: Marin Co.: Aedo (2000); Olema. June 7, 1936, Howell s.n. (NY): NOTES: This species, another relative of G. carolinianum L., otherwise occurs in east Texas and Louisiana (and the Azores). According to Aedo (2000), ““Its presence in California (not previously record- ed) probably constitutes a[n] occasional introduction’’. Hamamelidaceae Liquidambar styraciflua L.: DIST: ScV: CS: TEN: DOC: Sacramento Co.: North bank of American River, American River Parkway, a few hundred meters downriv- er of the Estates Dr. entrance. 38°33’N; 121°22’W. June 16, 1998, J. Randall s.n. (CDA, DAY). Hydrophyllaceae Wigandia caracasana HBK.: DIST: SCo: CS: NCI: DOC: Munz, PA. (1974, pg. 519): NOTES: No evidence for naturalization. Labeled specimens at CDA collected in the locations cited by Munz (1974), state that the plants were cultivated but without mention of spread from root- sprouts or other persistence mechanisms. Because this plant has an extensive root system it is a likely candidate to remain persistent from cultivation and spread locally. Hypericaceae Hypericum androsaemum L.: DIST: SnFrB: CS: N: DOC: Alameda Co.: Strawberry Creek behind UC- Berkeley Botanical Garden, June 16, 1993, B. Ertter 11898 (UC). Hypericum calycinum L.: DIST: CCo, SnFrB: CS: TEN: DOC: Contra Costa Co.: Berkeley-Oakland Hills, Grizzly Peak Boulevard ca. %4 miles south of Lomas Can- 86 MADRONO tadas junction, dense patch on east side of road at edge of mixed scrub, elev. ca. 100 ft. June 25, 1993, B. Ertter 11907 (UC); San Francisco Co.: Howell, J.T. et al. (1958, p. 102); Thomas, H. (1961, p. 239): NOTES: Gen. spread- ing vegetatively in California where it is widely planted and often persistent. Fully naturalized and spreading by seed in Oregon and Washington States, but similar behav- ior not confirmed in California. Hypericum hookerianum Wight & Arn.: DIST: NCo: CS: TEN: DOC: Mendocino Co.: forest road near Little North Fork Gualala River and Doty Creek, UTM Zone 10, 4298N, 4550E, elev. 100 ft, one large patch in second growth redwood forest, shrub 1—2 m tall. July 8, 2000, G. Leppig 1453 (CDA, HSC); Santa Barbara Co.: Monte- cito, Montecito School for Girls, escape from cultivation. June 9, 1951, H.M. Pollard s.n. (DAV, SBBG), det. by J.T. Howell. Lamiaceae Calamintha sylvatica Bromf. subsp. ascendens (Jor- dan) P.W. Ball: DIST: SnFrB: CS: TEN: DOC: Ala- meda Co.: Strawberry Canyon northeast of Panoramic Place in Oakland-Berkeley Hills, roadcut below oak for- est, small persisting patches at two sites along fire road. Sept. 2, 2000, B. Ertter 17518 (UC): NOTES: Nomen- clature as in Flora Europaea, which uses a narrow circum- scription of Satureja. If the same generic circumscriptions were applied in North America, no native Satureja would occur in California. An alternate name for this taxon is Satureja calamintha (L.) Scheele subsp. ascendens (Jor- dan) Brig. This is apparently the first record of the species occurring spontaneously in North America, although the closely related C. nepeta (L.) Savi is widely established in eastern North America. Cedronella canariensis (L.) Willd. ex Webb & Berth.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 119); Thomas, H. (1961, p. 299); Jep- son, W.L. (1943, pg. 400); Mud Lake, vicinity of San Francisco Bay. July 1914, A. King s.n. (JEPS). Galeopsis tetrahit L.: DIST: MP: CS: NCI: DOC: Mo- doc Co.: Corporation Ranch, Likely. July 8, 1958, 7.C. Fuller 1973 (CDA); Pit River Valley south of Alturas, irrigation ditch near S end of west side road, roadside. July 12, 1947, H.L. Mason and V. Grant 13417 (DAV, UC). Lamiastrum galeobdolon (L.) Ehrend. & Polatsch.: DIST: NCo: CS: TEN: DOC: Humboldt Co.: Arcata, Arcata Community Forest, 100 m S of trails 5 and 10 intersection. Second growth redwood forest. Large stolon- iferous mat. April 13, 2000, G. Leppig 1292, K. Neander (CDA, HSC): NOTES: Single 1/10 ha. patch. Represented here by a variegated cultivar, also sold as Lamium gal- eobdolon or Lamium variegatum. Lavandula stoechas L.: DIST: CCo, NCo: CS: C: DOC: Alameda Co.: sidewalk crack on Shattuck Ave. nr. Lincoln St. north Berkeley. June 25, 2001, B. Ertter 17699 (UC); Sonoma Co.: Fuller Mountain Rd., nr summit of Fuller Mtn., occasional on roadside in mixed conifer for- est. April 22, 2000, G. Leppig 1311 (CDA, HSC). Mentha xX villosa Huds.: DIST: SnFrB: CS: NCI: DOC: Napa Co.: Napa River riparian zone on gravel bar with willows, ca. % mile north of Trancas Street, city of Napa. El. 10 ft, TO6N, RO4W, Sec. 34, MD. Napa 7.5’ quad. Sept. 25, 1989, J. Ruygt 2408 (UC). Monarda citriodora Cerv.: DIST: ScV: CS: N: DOC: Sacramento Co.: roadside, Jackson Rd. (Hwy 16) W of Eagles Nest Rd. TO8N, RO6E, Sec. 25, MD. Elev. 125 ft. [Vol. 49 July 17, 1997, F. Carl s.n. (CDA); loc. cit. August 15, 1997, G.D. Barbe 4478 (CDA). Rosmarinus officinalis L.: DIST: CCo, SCo: CS: C: DOC: Alameda Co.: Albany waterfront, on former land- fill, several local shrubs. Aug. 19, 1994, B. Ertter 13077 (UC); Orange Co.: Newport Bay, North Star County Beach area on the SW end of the bay, ca. 2 km inland of Hwy 1, clearly spontaneous, not planted. Oct. 6, 1990, A.C. Sanders 10202 (DAV, UCR): NOTES: Single mature shrub also persisting near Huckleberry Regional Botanical Area, Alameda Co., spontaneous? Salvia longistyla Benth.: DIST: CCo: CS: N: DOC: Monterey Co.: Locally but abundantly spontaneous on bank of Big Sur River at Big Sur. Apr. 13, 1961, J.T. Howell 36487 (CAS, CDA); Munz, P.A. (1968, pg. 103); Howitt, B.E and J.T. Howell, (1973. pg. 29): NOTES: Cited on pg. 1343 of the Jepson Manual as “not natural- ized’”’; however field observation reports it extant and vig- orous in 2000, acc. to G. Norman via M.A. Matthews (personal communication to Hrusa, 4-2000). Salvia microphylla Benth.: DIST: CCo, NCoRO, SCo: CS: NCI: DOC: Marin Co.: Howell, J.T. (1970, pg. 358); Monterey Co.: Matthews, M.A. (1997, pg. 179).; Coal Chute Pt., dry sunny loam, originally cult. Aug. 14, 1936, L.B. Wheeler 4369. (Point Lobos State Reserve Herbari- um); Santa Barbara Co.: Munz, P.A. (1968, pg. 704); Sonoma Co.: Best, C., et al. (1996, pg. 164): NOTES: Cited on pg. 1343 of the Jepson Manual as “not natural- ized”’ under the name S. grahamii Benth. Salvia reflexa Hornem.: DIST: CaRE SNE: CS: GH/ C: DOC: Inyo Co.: garden in Independence, Rosedale Dr., elev. 4000 ft. July 24, 1996, M. DeDecker 6559 (CDA, RSA); Shasta Co.: garden of residence at 3657 Encanto Way northeast of Redding, surrounded by blue oak woodland. May 16, 1993, B. Ertter 11837 (UC). Salvia virgata Jacq.: DIST: MP, SBr., n SN: CS: NCI: DOC: Nevada Co.: just east of Grass Valley, on Empire Mine property. July, 1972, L. Mott. s.n. (JEPS); Empire Mine property, end of Stacy Lane off Highway 49, south side of Grass Valley, TI6N, RO8E, Sec. 34, MD. Weedy in 2-acre meadow. June 8, 1972, 7.C. Fuller and G.D. Barbe 964 (CDA, UC), det. by Ian C. Hedge (RGBE), Feb. 1987; San Bernardino Co.: Lake Arrowhead, garden escape. Aug., 1931, Braunton 1056 (DS); Siskiyou Co.: scattered plants on 600 sq. ft of drainage way in dry rangeland, adjacent to wet slough, Greenhorn Valley, ca. 2 mi W of Yreka. June 24, 1964, 7.C. Fuller 12244 (CDA), det. by E. McClintock; loc. cit. July 29, 1968, C.S. Giebner s.n. (CDA); Ager Beswick Road, very dry roadside. Aug., 1998, L. Parsons s.n. (SEPS): NOTES: Extirpated in Nevada Co; status of Siskiyou Co. plants currently under investigation. San Bernardino Co. speci- men originally identified as Salvia pratensis L. (sensu stricto). Acc. to R. Breckenridge (CDFA, Integrated Pest Control Branch), Salvia virgata is readily distinguished from S. pratensis s.s. by the foetid odor of its foliage. Scutellaria caerulea M. & S.: DIST: CCo: CS: C: DOC: Santa Clara Co: Weed in commercial field herb crop in Gilroy. Rocket Farms. Sept. 20, 1999, K. Meyer s.n. (CDA), det. by G.E Hrusa (UC). Stachys floridana Shuttlew.: DIST: ScV: CS: GH/C: DOC: Sacramento Co.: Abundant in garden, 2424 Park Estates Dr., Sacramento. May 21, 1963, K.S. Buchanan s.n. (CDA); loc. cit. June 7, 2000, L. Manger s.n. (CDA): NOTES: Tuberous perennial, spreading by rootsprouts. Present at this locality for at least 37 years. 2002] Lauraceae Cinnamomum camphora (L.) J. Presl: DIST: deltaic GV: CS: C: DOC: Contra Costa Co.: Antioch National Wildlife Refuge, Stamm Unit, two juvenile individuals on riparian margin, apparently spontaneous but source not evident. May 26, 2001, B. Ertter et al. 17563 (UC): NOTES: A fairly common landscape volunteer, but seed- lings generally restricted to irrigated sites and seldom al- lowed to mature. Laurus nobilis L.: DIST: CCo, NCo: CS: TEN: DOC: Humboldt Co.: Arcata, occasional in disturbed empty lot in redwood forest. Trees 2—4 m tall. April 12, 2000, G. Leppig 1289 (CDA, HSC); San Francisco Co.: Howell, leletedlen( los Saipo 7) hbomas: slies(l96L, spe. 173): NOTES: Single plant persisting on Albany Hill, Alameda Co., from unknown source and origin. Lentibulariaceae Utricularia subulata L.: DIST: NCoRO: CS: NCI: DOC: Mendocino Co.: Meyers-Rice (Madrono, in press); spreading through wet depressions and seeps in a pygmy forest, 39°15'’N; 123°45’W. Nov. 2, 1997, B. Meyers-Rice #MR971102 (DAV): NOTES: Intentionally introduced. Limnanthaceae Limnanthes macounii Trel.: DIST: CCo: CS: N: DOC: San Mateo Co.: Buxton (1998, pg. 184); Along east side of Hwy | south of Moss Beach, directly opposite Half Moon Bay airport, in cultivated field. Elev. ca. 10 m. Feb. 24, 2000, R. Schmid 2000-2 (UC): NOTES: Un- clear whether a locally naturalized alien or a previously overlooked native. Linaceae Linum trigynum L.: DIST: NCo: CS: NCI: DOC: Son- oma Co.: Best, C., et al. (1996, pg. 168). Malvaceae Anisodontea capensis (L.) Bates: DIST: NCoRO, ScV: CS: C: DOC: Sonoma Co.: Best, C., et al. (1996, pg. 171); Sacramento Co.: Among landscape shrubs along sidewalk in Sacramento. July 10, 2000, R. Gill s.n. (CDA): NOTES: In the nursery trade as Malvastrum ca- pense (L.) Gray & Harv. and so reported for Sonoma County (Best et al., 1996). Anoda pentaschista A. Gray: DIST: DSon: CS: NCI: DOC: Imperial Co.: Weed in citrus, nr Bard. Sept. 9, 1983, L. Pineda s.n. (CDA). Gossypium hirsutum L.: DIST: DSon, ScV: CS: NCI: DOC: Imperial Co.: Two ruderal plants betw. rd and base of canal bank, N side Hwy 98 to Mt. Signal 1.7 mi W of Calexico. Oct. 18, 1962, 7.C. Fuller 9804 (CDA); Sac- ramento Co.: City of Sacramento, Tahoe Park neighbor- hood, near the old fairgrounds, volunteer. Sept. 18, 1995, D. Goosen s.n. (DAV). Hoheria populnea A. Cunn.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Voluntary in dwarf conifer area, Strybing Arboretum, Golden Gate Park, San Fran- cisco. Sept. 20, 1973, G. Beutler s.n. (CDA). Lavatera olbia L.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Shrubs to 8 ft tall, commonly naturalized on non-irrigated waste ground of formerly cultivated gar- den, Stanyon St., San Francisco. Aug. 4, 1970, 7.C. Fuller se (EDA): Lavatera trimestris L.: DIST: SCo: CS: NCI: DOC: HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 87 Santa Barbara Co.: Smith, C.F (1976. pg. 192); Edge of water, Lauro Canyon Reservoir nr. San Roque Rd., Santa Barbara. June 25, 1975, C. Smith and J.L. Johnson s.n. (CDA). Malva verticillata L.: DIST: SCo: CS: NCI: DOC: Santa Barbara Co., Ventura Co.: Smith, C.F (1976, pg. LOS): Sida spinosa L.: DIST: SnJV: CS: NCI: DOC: Fresno Co.: near Sanger by the Ciba-Geigy Research Station on Annadale Ave., ca. 4% mi W of Reed Ave. Aug., 1996, B. Fischer s.n. (DAV). Moraceae Fatoua villosa (Thunb.) Nakai: DIST: SCo, SnJV: CS: GH/C: DOC: Kern Co.: Greenhouse weed, Arvin. Oct. 19, 1998, Lapp et al. s.n. (CDA); Riverside Co.: Sanders, A.C. (1996, pg. 527); San Bernardino Co.: Ontario. Weedy throughout nursery, under benches, in walkways. Oct. 21, 1983, Cohen s.n. (CDA); San Diego Co.: Nurs- ery containers, greenhouse, Fallbrook. Feb. 14, 1985, F. McCutcheon s.n. (CDA): NOTES: Also confirmed, but not vouchered, from Tehama Co., in commercial green- houses. Similar vegetatively to species of Laportea (Ur- ticaceae) and has been reported as that genus. Seed form readily distinguishes Fatoua from Laportea. Ficus palmata Forssk.: DIST: SCo: CS: NCI: DOC: Santa Barbara Co.: Spontaneous in creekbed, W Fk. Cold Spring Cyn., Santa Barbara. Dec. 23, 1958, H.M. Pollard s.n. (CAS, CDA): NOTES: Cited in Munz (1974) Smith, C.E (1976) and Rejmanek and Randall (1994) as F. pseudocarica Miq. Nymphaeaceae Nymphaea alba L.: DIST: NCo: CS: NCI: DOC: Men- docino Co.: Smith, G. and C. Wheeler (1990-1991, pg. 150). Oleaceae Fraxinus uhdei (Wenz.) Lingel.: DIST: SCo: CS: NW: DOC: Los Angeles Co.: ““Cottonwood Swamp’’, conflu- ence of San Francisquito Canyon stream and two tribu- taries draining off the southeast slopes of Red Mountain. Warm Springs Mtn. 7.5’ quad., TOSN, R16W, Sec. O01, elev. 1680-1690 ft [possible hybrid]. 7.S. Ross 7835 ((RSA? UCR, UC); San Bernardino Co.: Colton, S Pel- lisier Rd., near the corner of W Center St. and Orange St., S40 ily 20's W. LOZSROSW, SB? Bley, 262 m. Ephemeral creek with sandy bottom and the shaded grassy slopes above it. Nr. historical settlkement. Associated with Populus fremontii, Juglans californica and Rubus discol- or. Solitary sapling. Apr. 4, 1999, Mitch Provance 1763 (UCR): NOTES: Widespread and sometimes locally com- mon in coastal southern California riparian zones. For ex- ample, along the Santa Ana River near Riverside it forms a conspicuous element of the tree canopy near the conflu- ence of Spring Brook. Where street runoff flows into per- manently moist riparian areas, this species usually ap- pears. Identification is confused by the fact that it seems to hybridize with the F. velutina Torr. (F. pennsylvanica Marsh) complex, including both native plants and cv. ‘Modesto.’ Easiest to identify in winter because it is ev- ergreen, unlike the others. Much was cut last year in the area around Haskell Creek in the Sepulveda Basin during an effort to control exotics. Acc. to John Eckhoff (person- al communication to Sanders), they are “‘finding this tree in many of the riparian areas we visit or find ourselves 88 MADRONO working in, like San Gabriel River and Big Tujunga Wash at the base of the San Gabriel Mts.”’. Voucher requested but not received. Ligustrum lucidum W.T. Aiton: DIST: NCoRO, ScV, SnBr, SCo: CS: NW: DOC: Riverside Co.: Riverside, weed tree in landscaping on UCR campus, elev. 1100 ft. Dec. 16, 2000, A.C.Sanders 23728 (UCR); Sacramento Co.: Quinn, J., et al. (1991); San Bernardino Co.: San Bernardino Mtns., Thurman Flats Picnic Area, below Mountain Home Village, Yucaipa 7.5’ quad., TO1S, ROIW, Sec. 08, SB, 34°06'30"N; 117°00'05’W, elev. 3480 ft/1061 m, dense alder forest along the stream; solitary small tree ca. 3 m high in forest understory. May 27, 2000, A.C. Sanders 23432 (UCR): NOTES: This species is scat- tered around as an urban weed tree in the Riverside area; seeds apparently dispersed by birds. Also observed by Hrusa as seedlings and young trees in riparian zone along Arcade Cr., N side Interstate 80, Sacramento Co.; also by J. Ruygt (pers. comm., 3-2001), as seedlings on Redwood Rd., ca. 0.5 mi W of Hwy 29, and as seedlings and adult trees along bank of Camille Cr. At Polley Drive, both near Napa in Napa Co. Seldom collected, but apparently wide- spread in moist habitats. Ligustrum ovalifolium Hassk.: DIST: CCo, NCo, ScV: CS: NW: DOC: Mendocino Co.: Sinkyone Wilderness State Park, old homesite 2.7 miles south of Needle Rock Ranch House. Lost Creek trailhead in red alder woodland. June 25, 1995, F. Bowcutt 2009 (DAV, HSC); Rt. 1 near entrance to McKerricher State Park. Self-sustaining for over 50 years. July 30, 1981, G. L. Smith and C. R. Wheel- er 7205 (HSC). Monterey Co.: Elkhorn Slough National Estuarine Research Reserve, disturbed fields near South Marsh. May 22, 2000, G. Leppig 1382 (HSC); Sacra- mento Co.: Quinn, J., et al. (1991): NOTES: Known sites highly localized. Olea africana Mill.: DIST: SCo: CS: TEN: DOC: Riv- erside Co.: Riverside, Mt. Rubidoux, NE foot of the mountain above the end of 9th St., 33°59’N; 117°23'W, TO2S, RO2W, Sec. 22, SB. Elev. 1000 ft/305 m, E-facing decomposed granite slopes at edge between landscaped (residential) areas and coastal sage scrub. Disturbed and weedy with some no longer tended ornamentals. A locally common shrub or tree to 10 m tall. Oct. 23, 1996, A.C. Sanders 19643 (UCR): NOTES: Some individuals prob- ably originally planted (persisting ornamentals), but others growing in cracks of boulders, etc. and plainly spontane- ous. This is so scarce as a cultivated plant that this might easily be the only naturalized locality, where it is doing well. Reproduction is apparently by seed. Onagraceae Fuchsia magellanica Lam.: DIST: CCo, NCo: CS: NW: DOC: Contra Costa Co.: Cerrito Creek west of San Pablo Avenue, several reproducing shrubs on stream bank. Sept. 17, 1999, B. Ertter 16845 (UC); Humboldt Co.: Low shrub naturalized at base of Alnus sp., loop trail above Fern Canyon, Prairie Ck. Redwood State Park, 7.1 mi W of Highway 101 on Fern Canyon Rd. June 18, 1974, G.D. Barbe 1872 (CDA); Mendocino Co.: Mendocino, edge of headland behind Presbyterian church, uncommon shrub mixed in willow thicket. Sept. 29, 1992, B. Ertter 11449 (UC); Monterey Co.: Carmel Highlands, Fern Canyon east of Highway 1, shade of pine forest along creek. June 13, 1993, B. Ertter, V. Yadon, and M.A. Mat- thews 11890 (UC); Gibson Canyon, near Carmel High- lands. Growing in moist cyn. bottom, 6—10 pls. over sev- eral hundred yds, 600 ft elev. June 13, 1994, D. Kelch s.n. [Vol. 49 (DAV); San Francisco Co.: Lobos Creek between Lin- coln Blvd. and Baker Beach, deep shade. Nov. 15, 1992, B. Ertter 11459 (UC): NOTES: Some individual sites may be considered tenuous (TEN). Label data for second Mendocino Co. site in Sinkyone Wilderness State Park, [old home sites. T24N, R19W, Sec. 26, MD. May 23, 1989, F. Bowcutt 1325 (DAYV)], are not clear that location iS Spontaneous. Fuchsia X hybrida Voss.: DIST: SCo: CS: NCI: DOC: Ventura Co.: Persisting or spontaneous (?) in a clump of Ricinus shrubs on SPRR right of way, east Ventura. June 1, 1961, H.M. Pollard s.n. (CDA). Orobanchaceae Orobanche hederae Duby: DIST: CCo: CS: TEN: DOC: Alameda Co.: UC-Berkeley campus, small per- sisting colony in Hedera groundcover next to Koshland Hall. June 7, 2000, B. Ertter 17310 (UC); between student center and Alumni House. May 8, 2001, B. Ertter 17626 (UC). Papaveraceae Fumaria capreolata L.: DIST: CCo, SnFrB: CS: N: DOC: Alameda Co: Landscape weed in Livermore area. March 9, 1994, C. Elmore s.n. (DAV); Contra Costa Co.: Miller Knox Regional Park, at edge of excavation on west-facing hillside, localized but dense colony at base of coast live oak. Mar. 21, 1999, B. Ertter 16486 (UC); Ma- rin Co.: Muir Beach, 5.5 miles west of Hwy 1. Open area, slightly sandy soil, coastal strand with partial moisture, full sun. Not abundant. Elev. 50 ft. May 16, 1992, J.N. Le 17 (DAV); San Francisco Co.: Spontaneous along path betw. the Conservatory and Fuchsia garden, Golden Gate Park, San Francisco. Sept. 25, 1980, J.T. Howell 53901 (DAV); San Mateo Co: McClintock, E., et al. (1990, pg. 135): NOTES: Known sites widely scattered. Differing from the other naturalized species of Fumaria in California in the larger flowers (ca. 12 mm long) and broadly ovate sepals. Apparently becoming more com- mon. Similar to F. macrosepala Boiss. which may also be represented in California (Ertter 16486). Papaver X hybridum L.: DIST: CCo, ScV, SnJV: CS: NCI: DOC: Kern Co.: Point of Rocks, western (Kern) County. Apr. 27, 1950, E. McMillan and C. Smith 2684 (DAV); Antelope Valley, 3 mi W of Point of Rocks, the probable origin of the plants, not previously observed here by local farmers. Apr. 8, 1962, E. Twisselmann 6770 (CDA, DAV); Madera Co.: E side of Rd 26, 0.1 mi S of Ave. 12, 1 mi W of Madera, dominant weed on one acre of newly planted vineyard. Apr. 6, 1967, J.S. Davis s.n. (CDA); loc. cit. Apr. 19, 1967, T.C. Fuller 15609 (CDA, DAV); Sacramento Co.: SW corner of Metropolitan Air- port property near Garden Highway. Apr. 4, 1992, K. Mill- er s.n. (CDA, DAV); San Luis Obispo Co.: Choice Val- ley Hills, SE side of Sinsheimer Flat, dense colony in a bare area in a dry-farm wheat field (barley field in 7027). Apr. 29, 1962, E. Twisselmann 7025, 7027 (CDA, DAV): NOTES: The Twisselmann and J.S. Davis specimens were originally determined as Papaver apulum Ten. var. micranthum (Bor.) Fedde. Comment by Twisselmann in- dicated it is recent in San Luis Obispo Co. Passifloraceae Passiflora caerulea L.: DIST: NCoR, SnGb, SCo, s SN: CS: N: DOC: Los Angeles Co., Riverside Co. and San Bernardino Co:. Sanders, A.C. (Madrono, in press); 2002] Fresno Co.: Sequoia Mills, no date, K. Brandegee s.n., (UC); Napa Co.: Calistoga, July 11, 1910, K. Brandegee s.n. (UC): NOTES: Long persisting and difficult to erad- icate in garden situation, indicating strong potential to nat- uralize. Serves as host plant for non-native gulf fritillary caterpillars. Reported as non-spontaneous in Ventura Co. by H.M. Pollard (specimen at DAV, SBBG). Passiflora manicata (Juss.) Pers.: DIST: SCo: CS: NCI: DOC: Santa Barbara Co.: Smith, C.F (1976, pg. 197). Passiflora mixta L. f.: DIST: CCo: CS: N: DOC: San Francisco Co.: Golden Gate Park, local patch at junction of Crossover Drive and John EK Kennedy Drive. Sept. 11, 1993, B. Ertter 12269 (UC): NOTES: Passiflora mollis- sima auct. non (Kunth) L.H. Bailey as discussed under P. tarminiana. Freely reseeding in garden situation on Cat- alina Ave., Berkeley (Alameda Co.). Passiflora tarminiana Coppens & Barney: DIST: CCo: CS: N: DOC: Contra Costa Co.: edge of vacant lot at SW corner of San Pablo Avenue and Carlson Ave., climbing on Sambucus at edge of Cerrito Creek. Sept. 17, 1999, B. Ertter 16846 (UC): NOTES: Observed by Ertter to be also established upstream near BART path. Seed- lings occasionally encountered (and eliminated) by Ertter on Albany Hill, near Cerrito Creek. population; Reported also from Riverside Co. All original determinations as Passiflora mollissima auct. non (Kunth) L.H. Bailey, cur- rent determinations by D. Goldman. See Novon 11(1): pg. 9, 2001, for more information. Potentially a noxious pest, as in Hawaii (as P. mollissima sensu auct.), where capable of smothering native forests. Pedaliaceae Sesamum indicum L.: DIST: ScV, SNF: CS: C: DOC: Mariposa Co.: Few plants along roadside, Hwy 140 nr. Catheys Valley. Aug. 30, 1978, K.A. Parker s.n. (CDA): Sacramento Co: Single plant in asphalt divider, Kiefer Blvd, nr. Bradshaw Rd. Aug. 30, 1977, K. Miller s.n. (CDA): NOTES: Occurrences probably originating via commercial bird seed. Plumbaginaceae Limonium ramosissimum (Poir.) Maire subsp. prov- inciale (Pignatti) Pignatti: DIST: SCo: CS: NW: DOC: Santa Barbara Co., Ventura Co.: From Rick Burgess garden in Oxnard; originally collected in Carpenteria Salt Marsh, where it has naturalized about its mouth and is a very serious problem. 1994, Rick Burgess s.n. (SBBG): NOTES: Apparently from garden plants in an adjacent subdivision. Data provided by Dieter Wilken (SBBG). Polygalaceae Polygala myrtifolia L.: DIST: SCo: CS: NCI: DOC: Santa Barbara Co.: Smith, C.F (1976, pg. 183). Polygonaceae Polygonum multiflorum Thunb.: DIST: SnFrB: CS: GH/C: DOC: Marin Co.: Rampant weed in garden of Margadant Hayakawa, Eldridge Ave, Mill Valley. Vigor- ously spreading, but does not flower. Jan. 15, 1976, M. Hayakawa s.n (CDA), det. by E. McClintock, 9/1977: Cultivated in greenhouse [from roots dug at site of pre- vious collection], 3294 Meadowview Rd., Sacramento. Nov. 21, 1980, G.D. Barbe 3023 (CDA): NOTES: Dried tubers of this species (“Fo-ti” or ‘Ho Shou-wu’) are used as a folk remedy (Tyler 1982). HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 89 Polygonum orientale L.: DIST: SCo, ScV: CS: C: DOC: Sacramento Co.: spontaneous in residential yard, 2458 Catalina Dr. Sacramento. Aug. 11, 1975, F. Hine s.n. (CDA); Santa Barbara Co.: Smith, C.F (1976 pg. 122): NOTES: A specimen labeled only as “Mendocino Co., July 25, 1958, T. Erickson, Jr. s.n” is at CDA, but spontanaiety unknown. Other known occurrences are fu- gutives from cultivation. Ranunculaceae Caltha palustris L.: DIST: CCo, NCoR, NCo: CS: NW: DOC: Alameda Co.: Oakland Hills, Canyon east of Skyline Blvd, growing in stream that separates Huckle- berry Preserve from Sibley Volcano Park, along Skyline trail, nr. old homestead site. May 1, 1993, E.A. Dean 359 (DAV): Mendocino Co.: Smith, G. and C. Wheeler. (1990-1991, pg 151-152). Sonoma Co.: Rubtzoff. P. (1959, pp. 31-32): NOTES: Rubtzoff record also reported in Best, C., et al. (1996, pg. 201). Clematis terniflora DC.: DIST: ScV: CS: TEN: DOC: Sacramento Co.: Invasive in residental landscape and ad- jacent field on Larkspur Lane in Citrus Heights. Both sites relatively moist, both with sunny and shady situations. Climbing in Populus fremontii to 40+ ft, also in lower borders. Apr. 28, 2000, G.F. Hrusa 15389 (CDA): NOTES: Origin of infestation uncertain, but may have originated as garden ornamental. First noted as a pest in this site in 1992. Owners are attempting extirpation. Clematis vitalba L.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, pg. 72): Thomas, H. (1961, pg. 169): NOTES: Volunteer from planted ma- terial in Strybing Arboretum, Golden Gate Park, San Fran- cisco. Nigella damascena L.: DIST: SnFrB,. NCoRO: CS: N: DOC: Contra Costa Co.: waste ground, Brentwood Road at SPRR tracks, Brentwood. TOIN, RO2E, Sec. 18, MD. June 18, 1974, J. deFremery and C. Butler s.n. (CDA): Sonoma Co: Best, C., et al. (1996, pg. 203): NOTES: Garden escape, naturalized as a weed about habitations. Ranunculus cortusifolius L.: DIST: CCo: CS: TEN: DOC: Alameda Co.: Berkeley, University of California campus, untended garden plots at NW corner of Valley Life Sciences Building. May 8, 2001, B. Ertter 17625 (UC): NOTES: Possibly originating from deliberately strewn seeds, but now self-sustraining and spreading to adjacent untended plots. Determination by R. Ornduff, 2000. Rhamnaceae Ziziphus jujuba L.: DIST: ScV: CS: C: DOC: Sacra- mento Co.: Ditch at corner Sheldon Rd and Hwy 99, sap- ling. July 7, 1998, N. Wymer s.n. (CDA): Yolo Co.: 15— 20 plants up to 1 meter tall on W side of Hwy 113 appr. 1 mi N of Covell exit in Davis. Obviously spontaneous, some inside ROW. Aug. 21, 2001, D. Adams s.n. (CDA): NOTES: Escape from cultivation, perhaps from garden trash. Rosaceae Cotoneaster lacteus W.W. Smith: DIST: CCo, SnFrB: CS: NW: DOC: Alameda Co.: Albany Hill, scattered col- ony. Feb. 9, 1997, Ertter 15449 (UC): Contra Costa Co.: Ygnacia Valley Road cut through Lime Ridge, single shrub. Jan. 5, 1997, Ertter 15432 (JEPS): San Mateo Co.: McClintock, E., et al. (1990, pg. 144): NOTES: Noted elsewhere in the east San Francisco Bay Area. Flowers 90 MADRONO like C. pannosus but leaves larger, 3.5—7 cm long, obvi- ously depressed-veiny, obovate-elliptic with obtuse apex; inflorescence often more floriferous as well. Crataegus monogyna Jacquin: DIST: NCoRO, SnFrB: CS: NW: DOC: Alameda Co.: Dry Creek/Pioneer Re- gional Park east of Hayward. Jan. 19, 1992, B. Ertter 10894 (UC); Anthony Chabot Regional Park, Cascade Trail. April 29, 1981, Jack Stratford s.n. (JEPS); also ob- served by Ertter as well-established in Sibley Volcanic Regional Preserve, Oakland Hills; San Mateo Co.: McClintock, E., et al. (1990, pg. 102): NOTES: Also ob- served as widely scattered plants on the Palisades, south shoulder of Mt. St. Helena in Napa Co. Thorny shrub to small tree, leaves 3—5-lobed halfway or more to midvein. Cydonia oblonga Mill.: DIST: ScV, SCo, SnJV: CS: NCI: DOC: Sacramento Co.: bush 10 ft tall and across, one of a number of plants persistant from cult. in a fence- row, W side of Elk Grove-Florin Rd., 0.1 mi N of Sheldon Rd., Elk Grove. Nov. 19, 1969, 7.C. Fuller 19101 (CDA); Santa Barbara Co.: Smith, C.E (1976, p. 160); Stanis- laus Co.: % mi W of La Grange, Nof Hwy 132, woodland at edge of dredge tailings, occasional escape in this area. April 3, 1969, P. Allen 123 (DAV): NOTES: Sacramento County site now in developed area, probably extirpated. Cydonia sinensis Thouin: DIST: NCo: CS: NCI: DOC: Mendocino Co.: Smith, G. and C. Wheeler, (1990— 1991, pg. 181): NOTES: Reported as Chaenomeles si- nensis (Thouin) Koehne. Eriobotrya japonica Lindl.: DIST: SnFrB, SCo: CS: NCI: DOC: Alameda Co.: Oakland, Joaquin Miller Park, Sunset Trail. Occasional in redwood forest. Dec. 4, 2000, G. Leppig 1521 (HSC); Santa Barbara Co.: Smith, C.F (1976, p. 160): NOTES: Seedling observed on Albany Hill (Alameda Co.) by Ertter. Expected elsewhere. Filipendula vulgaris Moench: DIST: SnFrB: CS: NCI: DOC: Alameda Co.: few scattered plants uphill from Australasian beds, Univ. of California Botanic Garden, Strawberry Canyon, Berkeley. Oct. 12, 1978, G.D. Barbe 2459 (CDA). Photinia davidsoniae Rehd. & Wilson: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 84): NOTES: Probably no more than a casual escape from cultivation. Potentilla anglica WLaicharding: DIST: NCo, SCo, ScV, SnFrB: CS: NW: DOC: Contra Costa Co.: Ertter, B. (1997, pg. 78); Bishop Ranch Regional Open Space southwest of San Ramon, shaded streambed. Aug. 8, 1993, B. Ertter and B. Olson 12198 (UC); Mendocino Co.: Smith, G. and C. Wheeler, (1990-1991, pg. 184); Sacramento Co.: Wet seep behind house, S end of Blue Oak Flat at summit of American River Bluffs, Folsom Lake State Park, Natoma Unit. Perhaps not persisting. May 21, 1990, G.F. Hrusa 7930 (CDA); San Diego Co.: Poway, Blue Sky Ecological Reserve, Oak Grove area of park, stream edge. Aug. 22, 1995, M. Hanson s.n. (UC): NOTES: Included in Munz and Keck (1959) but left out of Jepson Manual due to lack of sufficient evidence avail- able at the time that this species was truly naturalized in California. Potentilla reptans L.: DIST: SnJV: CS: TEN: DOC: Fresno Co.: Town of Fresno, a lawn weed. June, 1997, B. Fischer s.n. (CDA, DAV). Prunus laurocerasus L.: DIST: NCo, SCo: CS: NW: DOC: Humboldt Co.; Arcata, occasional, naturalized in redwood forest behind Humboldt State University, west of Fern Lake, April 12, 2000, G. Leppig 1291 (CDA, HSC); Santa Barbara Co.: upper Alturas Rd., Santa Barbara. [Vol. 49 Sept. 30, 1939, M. Van Rensselaer 1343 (UC): NOTES: Escaping from Humboldt State University landscaping. Prunus persica (L.) Batsch: DIST: CaR, SnFrB: CS: C: DOC: Contra Costa Co.: south base of Mount Diablo, lone tree in Sycamore Canyon, Apr. 25, 1995, B. Ertter and C. Thayer s.n. (JEPS); Los Angeles Co.: Whittier Hills, (Puente Hills, pro parte): Turnbull Canyon nr. mouth. Three trees, escape or persistent from cult.?, no habitations nearby. T2S, R11W Sec. 22. Elev. 505 ft. Apt. 21, 1991, T. Ross 4499 (CDA, RSA, UC); Siskiyou Co.: Mount Shasta, along Everett Memorial Hwy, adventive in chaparral. Elev. 4300 ft. July 4, 1968, W.B. and V.G. Cooke 39264 (UC). Prunus serrulata Lindl.: DIST: SnFrB: CS: C: DOC: Alameda Co.: Strawberry Canyon firetrail behind UC Berkeley Botanical Garden, several small trees in shady wooded area, Apr. 15, 1999, B. Ertter 16502 (UC). Pyracantha coccinea M. Roem.: DIST: NCoRO, SnFrB: CS: TEN: DOC: Alameda Co.: Strawberry Can- yon, at Poultry Farm Station, 500 ft, Oct. 6, 1934, C.M. Belshaw 218 (UC). Sonoma Co.: Best, C., et al. (1996, jog, ZS). Pyracantha crenatoserrata (Hance) Rehder: DIST: CCo, GV, SCo, SnFrB: CS: N: DOC: Contra Costa Co.: Lime Ridge open space, occasional shrub in grassland of quarry area, May 10, 1998, B. Ertter 16115 (JEPS); Sac- ramento Co.: Moist disturbed areas in vicinity of Willow Creek parking area, E side Lake Natoma. Common. 38°37'N; 121°02'W. Elev. 75 m. May 2, 1990, G.F. Hrusa 7802 (CDA); loc. cit. July 26, 1990, G.F. Hrusa 8155 (CDA); San Mateo Co.: McClintock, E., et al. (1990, pg. 147); Ventura Co.: Seedlings growing in railroad gravel ballast at Foster Park, Ventura. Aug. 14, 1964, H.M. Pol- lard s.n. (CAS, CDA); Escape from cultivation in brush strip under Ventura River bluff N of and nr Santa Ana Blvd. crossing, Oak View. Oct. 23, 1963, H.M. Pollard s.n. (CAS, CDA): NOTES: Distinguished by the narrowly obovate leaves to 2 cm wide, often marginally toothed and apically notched. Also observed by Ertter on Albany Hill in Alameda Co. Common as seedlings in urban gardens, lawns. Expected elsewhere. Pyracantha crenulata (D. Don) M. Roem.: DIST: NCoRO, SnFrB: CS: TEN: DOC: Sonoma Co.: Best, C., et al. (1996, pg. 215): NOTES: Also observed by Ertter on Albany Hill, Alameda Co. Pyrus communis L.: DIST: SCo, ScV, SNE, SnFrB: CS: N: DOC: Contra Costa Co.: EBMUD land east of Tilden Park, ca. % mile southeast of Inspiration Point, grazed grassland near streamlet, single well browsed large shrub. Possibly only persisting from cultivation. May 7, 1993, B. Ertter and C. Thayer 11700 (UC); Mono Co.: Spring 0.7 mile S of the mouth of Milner Creek, 0.6 mile N 08° W of Copper Queen Mine, TO4S, R33E, Sec. 21, MD. Elev. 5210 ft. Persisting apparently several decades after introduction, compact trees ca. 4 m high in silty moist alluvial soil with Chrysothamnus, Stanleya, Robi- nia. May 29, 1986, James D. Morefield and Douglas H. McCarty 3718 (RSA, UC); Orange Co.: Santa Ana Mountains, 4 mi NE Lake Irvine, Black Star Canyon, Cleveland National Forest, Trabuco District. Oak wood- land canyon with riparian woods along the stream. Grassy clearings and chaparral on the canyon slopes. April 18, 1966, E.W. Lathrop 6176 (RSA); Southern Santa Ana Mountains, San Mateo Canyon Wilderness Area. Western edge of the wilderness in Lucas Canyon along canyon floor from crossing of Lucas Canyon Trail, upstream % mile to spur trail leading to old mining habitations in side canyon, T07S, RO6W, Sec. 10, SB. Elev. 1000—1350 ft. 2002] Low shrub, single individual, apparently adventive about old homestead. May 20, 1992, S. Boyd and T. Ross 7468 (RSA); Santa Barbara Co.: Smith, C.F (1976, pg. 164); Santa Clara Co.: Coyote Creek. Mar. 30, 1901, A.P. Chandler 911 (UC); Sonoma Co.: Pitkin Marsh, near Fo- restville, eastern branch of the Upper Marsh. At the edge of a thicket, on damp ground. Oct. 26, 1952, P. Rubtzoff 1332 (CAS, RSA); W branch upper marsh. Oct. 26, 1952, Rubtzoff 1329 (UC); Ventura Co.: Matilija Canyon, off- spring of old ranch planting in creek bottom. Oct. 6, 1946, H.M. Pollard s.n. (RSA, SBBG); Munz, P.A. (1974, pg. 758): NOTES: Sacramento Co.: observed as a possible escapes adjacent to orchards approx. one mile S of Free- port. Rosa multiflora Thunb. ex Murray: DIST: CaR, CCo, ScV: CS: NCI: DOC: Butte Co.: ca. 5 airmiles southwest of Gridley, south side of Evans Reimer Rd. ca. % mile east of Pennington Road, T17N, RO2E, Sec. 07, MD. 70 ft, May 2, 1998, L. Ahart 7958 (JEPS): San Mateo Co.: McClintock, E., et al. (1990, pg. 148): Siskiyou Co.: Mt. Shasta City, sedge meadow at corner of Lake St. and Commercial St. June 9, 2001, B. Ertter 17662 (UC): Shas- ta Co.: Squaw Creek Canyon, April 19, 1992, D.W. Tay- lor 12602 (UC). Det. by B. Ertter, 11-2001. Rubus ulmifolius Schott var. ulmifolius: DIST: CCo, NCoR: CS: N: DOC: Napa/Lake Co.: Highway 29 on north side of Mount St. Helena, roadside at edge of ser- pentine area, locally common, forming large tangles. Nov. 27, 1997, B. Ertter and L. Constance 15885 (UC): NOTES: Also observed in the Presidio of San Francisco (San Francisco Co.) by Ertter. Rubiaceae Coprosma repens A. Rich.: DIST: CCo: CS: N: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 130); Thomas, H. (1961, p. 323); San Mateo Co.: McClintock, E., et al. (1990, pg. 149): NOTES: Observed in Monterey Co. by Sanders, (10-2000) on seacliffs in Pacific Grove. An escape from cultivation. Salicaceae Populus nigra L. cv. ‘Italica’: DIST: SCo, SNE, NCoRO: CS: TEN: DOC: Alameda Co.: Tree NW of Botanical Garden, Strawberry Canyon, Berkeley. July 10, 1943, B. Rodin 245 (DAV): Sonoma Co.: Best, C., et al. (1996, pg. 221); Mono Co.: mouth of Milner Creek, spring 0.7 mi S; 0.6 mi n 83 degrees w of Copper Queen Mine, Owens Valley drainage, White Mountains. Silty moist alluvial soil sloping 3% SW _ w/Chrysothamnus, Stanleya, Salix, Rosa, Artemisia. Elev. 5210 ft. T04S, R33E, Sec. 21, MD. May 29, 1986, James D. Morefield and Douglas H. McCarty 37l6e (UC): Ventura Co.: Spreading by root-sprouts from parent tree along ditch on Loma Dr. S of Ventura Ave, Ojai Valley, Ventura. Nov. 8, 1967, H.M. Pollard s.n. (CAS, CDA): NOTES: Per- haps original plants persistent from cultivation, but often spreading aggressively by root sprouts. Best et al. (1996) report that only staminate plants are in cultivation in Cal- ifornia. Sarraceniaceae Sarracenia aff. rubra Walter: DIST: NCo: CS: N: DOC: Mendocino Co.: Smith, G. and C. Wheeler (1990— 1991, pg. 171): NOTES: Intentionally introduced. Ac- cording to B. Meyers-Rice (DAV) the pitcher plants intro- duced in this site are represented by hybrid forms. No HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 91 clear S. rubra has been as yet found, although some in- dividuals may outwardly approach it in appearance. Scrophulariaceae Anarrhinum bellidifolium (L.) Willd.: DIST: SnFrB: CS: C: DOC: Alameda Co.: Strawberry Canyon firetrail behind UC Berkeley, 500 m elev., July 18, 1999, N. Hill- yard s.n. (JEPS): NOTES: Determination verified by B. Ertter and M. Wetherwax. Limnophila X ludoviciana Thieret: DIST: ScV: CS: TEN: DOC: Butte Co.: Rice field at NE corner of Ham- ilton and Riceton Rds, Rice Experiment Station NW of Biggs. Scattered plants on moist mud betw. experimental rice plots. TION, RO2E, Sec. 35, MD. Elev. ca. 100 ft. Oct. 22, 1998, Oswald, Ahart, Ertter 9674 (CDA, CHSC): Rice Experiment Station near Biggs, mud of drained ex- perimental plots, rare. Oct. 22, 1998, B. Ertter 16446 with L. Ahart and V. Oswald (UC), det. by D. Philcox (K), 1998; Yuba Co.: Ahart, L. (1981, pgs. 7, 8): In shallow water in open area in a rice field, 2 miles NE of Marys- ville. Sept. 25, 1977, L. Ahart s.n. (CAS, CDA, CHSC) det. by D. Philcox (K), Dec., 1977: NOTES: =Limno- phila indica (L.) Druce X L. sessiliflora Blume. Reported by Ahart (1981) as L. indica. Examination of both Yuba and Butte Co. material by Philcox (K) indicates our plants are best referred to the hybrid form. Weed of agriculture, subject to elimination via drainage of cultivated rice fields. Mazus japonicus Kuntze: DIST: SCo: CS: GH/C: DOC: Orange Co.: Bordier’s Nursery, Irvine Boulevard, Santa Ana, spontaneous in locally prepared soil mix. Aug. 9, 1973, G.D. Barbe 1694 (CDA, UC). Penstemon strictus Benth.: DIST: SNE: CS: TEN: DOC: Mono Co.: Witcher Creek, at crossing of jeep road (Inyo National Forest road 4854) from Swall Meadows. June 29, 1988, D.W. Taylor 9917b (JEPS): NOTES: Ap- parently escaping from nearby rural residential areas of Swall Meadows, either from cultivation or as seeded for ‘wildflowers’; common in unmanaged ruderal vegetation of vacant lots in the developed portion of the subdivision. The Witcher Creek location is in an area of natural veg- etation removed some 500 m and over a small hill from the most proximal habitations. The occurrence was still present in 1998. Report and data provided by D.W. Taylor. Penstemon subglaber Rydb.: DIST: SNE: CS: TEN: DOC: Mono Co.: Mammoth, vacant lot near the Post Of- fice. Aug. 2, 1998, D.W. Taylor 16939 (JEPS): NOTES: Apparently escaping from nearby areas seeded for ‘wild- flowers.’ The plants reseed in unmanaged, ruderal vege- tation in the developed portion of town, but have not yet been seen afar. Report and data provided by D.W. Taylor. Scrophularia peregrina L.: DIST: SCo: CS: N: DOC: Los Angeles Co.: Ross, T. and S. Boyd (1996, pg. 436): Claremont, RSABG; SE edge of Indian Hill Mesa, ver- nally moist clay embankment. Common winter and spring weed in the area. March 24, 1993, T. Ross 6819 (CDA, RSA, UC): NOTES: Introduced in 1950s during biosys- tematic study of Scrophularia. Robust specimens may mimic Scrophularia californica. Native to Mediterranean region. Verbascum olympicum Boiss. non Bunyard: DIST: NCoRI: CS: NW: DOC: Sonoma Co.: Common on benches above rocky, dry bed of Leslie Creek from approx. Mark West Ck. confluence to first main tributary from NE. Mixed with Verbascum thapsus. July 25, 2000, G.F. Hrusa 15690, 15691, 15692 (CDA, and to be dis- tributed): NOTES: Readily distinguished from other nat- uralized Verbascum by its dense white-arachnoid pubes- 92 MADRONO cence, especially in the inflorescence. Flowers are brighter yellow and larger than in V. thapsus. First escaping from ornamental plantings at residence on CAS Pepperwood Preserve in 1976. Population, mixed with Verbascum thapsus, is large and apparently increasing. Possible hy- brid forms are also scattered at this locality (July 25, 2000, Hrusa 15693, CDA). Solanaceae Atropa belladonna L.: DIST: CCo, ScV: CS: NCI: DOC: Sacramento Co.: Hampton Rd., Sacramento. Gar- den weed. Aug. 24, 1999, Ken and Mary Brown s.n. (CDA); San Francisco Co.: Howell, J.T. et al. (1958, p. 121); Thomas, H. (1961, p. 304). Capsicum annuum L.: DIST: SCo: CS: NCI: DOC: Ventura Co.: Spontaneous in fallow field on lower Pier- pont Bay, Ventura. Sept. 15, 1959, H.M. Pollard s.n. (CDA). Cestrum parqui L’Her.: DIST: c SNE SCo, SnFrB: CS: NCI: DOC: Amador Co.: S side of Jackson on rd to Mokelumne Hill. Aug. 9, 1933, C.B. Wolf 5206 (UC); Napa Co.: Napa City. 1893, no collector (UC); Santa Barbara Co.: Toro Canyon Creek at Southern Pacific RR crossing, Summerland. Aug. 8, 1961, H.M. Pollard s.n. (CDA, SBBG). Lycium ferocissimum Meirs: DIST: SCo: CS: NCI: DOC: Los Angeles Co.: shrub 3 m tall, on canal bank in salt marsh, 400 m N of 431 East Culver Blvd, Playa del Rey. TO2S, R1I5W, Sec. 27, SB. Jan. 16, 1979, T.C. Fuller 20255 (CDA): NOTES: Persistent from cultivation? Nicotiana X sanderae Hort. ex Wats.: DIST: CCo: CS: C: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 121); Thomas, H. (1961, p. 305). Nicotiana tabacum L.: DIST: SnFrB, SCo: CS: C: DOC: Contra Costa Co.: | volunteer plant at waterfront at E end of building, Golden Gate Fish Co., Point San Pablo, Richmond. Dec. 20, 1966, R. DeBoer s.n. (CDA), det. by T.-C. Fuller; San Diego Co.: Cholla nr. San Diego. Apr. 7, 1885, Cleveland and Greene s.n. (UC); Santa Barbara Co.: Smith, C.F (1976, pg. 251). Petunia violacea Lindl.: DIST: CCo, SCo: CS: NCI: DOC: San Bernardino Co.: Alta Loma, Hellman and 19th St., roadside. April 2, 1961, G. Pilone 140 (DAV); San Luis Obispo Co.: roadside, no dwelling in vicinity, Perfumo Canyon Rd, 0.9 mi W of junct. with Los Osos Valley Rd., ca. 5 mi SW of San Luis Obispo. July 19, 1962, 7.C. Fuller 9237 (CDA); Ventura Co.: Naturalized locally along San Antonio Creek at Royal Oaks Dairy and for some distance downstream, Ojai. Aug. 13, 1966, H.M. Pollard s.n. (CDA, SBBG): NOTES: An escape from cul- tivation but extent of true naturalization not known. Solanum gayanum (Remy) Phil. f.: DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 122); Thomas, H. (1961, p. 303): NOTES: Escape from cultivation. Solanum scabrum Mill.: DIST: ScV, SnJV: CS: C: DOC: Butte Co.: Six volunteer plants, city dump, Sterling City. T24N, RO4E, Sec. 28, MD. Sept. 11, 1974, Sauer and Heinricks s.n. (CDA), det. by T.C. Fuller; Fresno Co.: Town of Five Points, growing at the Westside Research and Extention Center in mix of seeds of Solanum nigrum complex planted in tomato field for herbicide trial. 1999, Bill Fisher s.n. (DAV); Sacramento Co.: Vigorous growth to 8 dm, vacant lot W side of 1226 D St. Sacra- mento. Aug. 26, 1988, G.D. Barbe 4310 (CDA). Solanum villosum Mill.: DIST: ScV: CS: C: DOC: [Vol. 49 Yolo Co.: Knights Landing, tomato field near wet area. May, 1996, G. Miyou s.n. (DAV). Urticaceae Boehmeria cylindrica (L.) Sw.: DIST: GV: CS: NW: DOC: Sacramento Co.: Hrusa, G.E (2000, pgs. 138-— 139); NOTES: Observed by Hrusa as becoming increas- ingly common about Snodgrass Slough, Sept. 2001; also observed by Hrusa as common in Sutter Slough and Elk Slough (Yolo Co.) immed. W of Courtland, Aug. 2002. Urtica-like foliage but without stinging hairs, the opposite leaves and inflorescence of sessile, aggregate clusters su- perficially resemble Lamiaceae. Readily distinguished by the mostly ebracteate inflorescence (although leafy at the apex), scabrous leaves, round stems, absence of a corolla, and unisexual flowers. Boehmeria cylindrica may be mon- oecious or dioecious, but all California material examined to this point is monoecious, with staminate flowers fewer and confined to the uppermost parts of the spike. Laportea aestuans (L.) Chew: DIST: SCo: CS: GH/C: DOC: San Diego Co.: Weed in greenhouses. Commercial nursery, Fallbrook. April, 1999, A. Amador s.n. (CDA): NOTES: Vegetatively similar to Fatoua villosa (TYhunb.) Nakai, (Moraceae). Seed form unequivocally distinguishes F. villosa from the genus Laportea. Verbenaceae Verbena rigida Spreng.: DIST: CCo, NCoR, ScV, SnFrB: CS: NCI: DOC: Contra Costa Co: few scattered plants in lawn, Elmwood Rd, El Sobrante. Sept. 3, 1965, R. DeBoer s.n. (CDA); Marin Co.: Spontaneous in filled ground, San Pedro Rd. at Marin Yacht Club, San Rafael. Filled ground, edge of Yacht Harbor. Sept. 23, 1965, Gor- don True 2652 (CDA); Sacramento Co.: spontaneous in sidewalk, 15th and P Sts., Sacramento. Aug. 15, 1961, R.M. Hawthorne s.n. (CDA); Sonoma Co.: Howell J.T. (1972, pg. 102): NOTES: Sometimes treated as V. venosa Gillies and Hooker. Vitex agnus-castus L.: DIST: SCo, ScV: CS: NCI: DOC: Santa Barbara Co.: naturalized on SPRR tracks at Ortega St., Santa Barbara. June 11, 1968, P. Okuye s.n. (CDA); Yolo Co.: Creek by apiary, University Farm Cam- pus. Oct. 14, 1932. H.A. Barthwick s.n. (DAV). Vitaceae Cissus antarctica Venten.: DIST: SCo, SnFrB: CS: NCI: DOC: Alameda Co.: Escaped ornamental, flood channel, Union City. Feb. 2, 1981, Sweigert s.n. (CDA); Los Angeles Co.: Weed in CalTrans yard betw 710 and Ist. St., East Los Angeles. Oct. 9, 2001, M. Adams s.n. (CDA, UCR). Vitis aestivalis Michx.: DIST: SCo: CS: NCI: DOC: Ventura Co.: Apparently spontaneous, Valley Rd nr. San- ta Ana Blvd., Oak View. June 2, 1972, H.M. Pollard s.n. (CAS, CDA). Vitis rupestris Scheele: DIST: NCoRO: CS: TEN: DOC: Sonoma Co.: Best, C., et al. (1996, pg. 247): NOTES: Winegrape rootstock. Zy gophyllaceae Peganum harmala L.: DIST: DMoj: CS: EXT?: DOC: Kern Co.: Edwards AFB, roadside % mi E of Lancaster Blvd, on Old Hospital Rd. Across from “‘P”’ housing sec- tion. Single plant. TOON, R1OW, Sec. 14, SB. June 29, 1990, D. Charlton 4453 (CDA); San Bernardino Co.: Abandoned 15 acre pasture, Minneola and Elkhorn Rds, 2002] Newberry Springs. TOON, RO2E, Sec. 27, SB. July 8, 1988, J. Hitchcock s.n. (CDA); 50-60 clumps, 3-5 ft diam. in abandoned pasture, Minneola and Elkhorn Rds, Daggett. T11N, ROLE, Sec. 27, SB. July 1, 1988, J. Hitch- cock s.n. (CDA); Newberry Springs, vicinity of Silver Valley Rd. N of RR tracks nr the airport, in yards of res- idents and vacant lots. Aug. 23, 1988, D. Pendleton s.n. (DAV): NOTES: Noxious Weed under eradication by Cal- ifornia Dept. of Food and Agriculture. ANGIOSPERMS—-MONOCOTS Alismataceae Sagittaria brevirostra Mackenzie & Bush: DIST: CCo: CS: NCI: DOC: Marin Co: Chileno Valley, the laguna. June 18, 1947, J.T. Howell 23261 (UC): NOTES: Specimen originally determined as S. latifolia; current de- termination by K. Rataj, 1968, with confirmation by C.B. Hellquist, 1994. Cited in Haynes and Hellquist (2000). Sagittaria rigida Pursh: DIST: CaRH, CCo, NCoRI: CS: NW: DOC: Oswald, V.A. et al. (1998, p. 185): NOTES: Introduced to ‘improve/enhance’ waterfowl hab- itat? Araceae Arum palestinum Boiss.: DIST: NCo: CS: TEN: DOC: Humboldt Co.: Arcata Marsh and Wildlife Sanctuary, Butchers Slough upper salt marsh. May 8, 1998, G. Lep- pig 755 (CDA, HSC); Humboldt Bay, Woodley Island near hunting cabin, moist Juncus meadow. UTM Zone 10 4518N 4020E. July 14, 1976, Peter Sorenson s.n. (HSC): NOTES: This species, although locally present in small numbers is long-lived and thus highly persistent. Arcata Marsh population under eradication by City of Arcata. Dracunculus vulgaris Schott: DIST: SCo, SnFrB: CS: N: DOC: Alameda Co.: ca. halfway up Claremont Can- yon in Oakland Hills, locally established colony on steep roadfill. June 16, 1995, B. Ertter and L. Constance 14258 (UC); loc. cit. June 7, 1992, B. Ertter 11082 (UC): Santa Barbara Co.: Santa Barbara, growing in lot probably once under cultivation. May 17, 1948, R.S. Beal, Jr. s.n. (UC): NOTES: Claremont Canyon colony still present as of April 1990, possibly expanding. Pinellia ternata (Thunberg) Makino: DIST: SnFrB: CS: NCI: DOC: San Francisco Co.: Golden Gate Park, “accidentally introduced”’. May, 1935, E. Walther s.n. (UC): NOTES: Determined by S.A. Thompson, 1994; original determination as P. tripartita (Blume) Schott. Cyperaceae Bulbostylis barbata Kunth: DIST: CCo: CS: GH/C: DOC: Santa Cruz Co: Weed growing in commercial greenhouses, Watsonville. Probably originating from Sri Lanka. May 14, 1998, T. Watson s.n. (CDA). Cyperus flavescens L.: DIST: ScV: CS: NW: DOC: Butte Co.: damp sand, E side Sac. R., NW Parrott Land- ing, 1 mi SE Ord Ferry, 12 mi SE Chico. Aug. 13, 1999, Lowell Ahart and V. Oswald 8143 (CDA, CHSC), det. by V. Oswald, 10-99; loc. cit. Oct. 28, 1999, L. Ahart 8303 (CHSC): Arch Rock tunnel, Feather River Hwy, Elev. 500 m. Sept. 6, 1981, L. Ahart 3123 (CHSC, DAV): Peter Ahart Ranch, 1 mi N and 2% mi E of Honcut. July 19, 1975, L. Ahart 901 (CHSC); loc. cit. Aug. 21, 1975, L. Ahart 957 (CHSC); loc. cit. Sept. 4, 1987, L. Ahart 5877 (CHSC); loc. cit. Aug. 19, 1995, L. Ahart 7618 (CHSC, JEPS); wet sand on N. Fk. Feather River nr Poe Power- house, riparian woodland, elev. 890 ft. Aug. 30, 1987, L. HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 93 Ahart 5865 (CHSC); wet sand on margins of small pond in Oroville Wildlife Area. Common. Elev. 140 ft. Aug. 23, 2000, L. Ahart 8664 (CHSC): moist gravel bar along Sacramento R. 0.25 mi downstream from Murphy’s Slough. Riparian woodland. July 31, 1983, V. Oswald 94] (CHSC); Feenstra’s Riverview Orchard W end of Cana Hwy between S half Cana Lake (on Dicus Slough) and Sacramento River. Uncommon. Wet gravel near shallow pools on large gravel bar along river. Portion of Rancho Bosquejo, elev. 160 ft. T23N, RO2W, MD. Aug. 13, 1987, V. Oswald 3215 (CHSC): Tehama Co.: Hog Lake Plateau along Hwy. 36 NE of Red Bluff, T28N, RO3W, Sec. 14, MD, elev. 430 ft, moist soil along Paynes Creek ca. 1 mi upstream from the old bridge site. Sept. 4, 1996, V. Os- wald and L. Ahart 8270 (CHSC), det. L. Janeway; Yuba Co.: Wet soil in irrigated pasture on Eugene Ahart Ranch in Marysville, elev. 100 ft. Sept. 11, 1975, L. Ahart s.n. (CHSC), det. V. Oswald; shallow water on disturbed rocky soil, S side Scott Forbes Rd, 0.25 mi E of intersection Scott Forbes Rd and Peoria Rd. E of Gray Dr. E of Browns Valley. Uncommon, el. 300 ft. Aug. 13, 1997, L. Ahart 7865 (CHSC), det. V. Oswald: NOTES: Additional collections from some of these same localities are at CHSC, the earliest from 1961. Cyperus flavicomus Michx.: DIST: ScV, SNF: CS: N: DOC: Butte Co.: University of California Biggs Rice Re- search Station, near the corner of Riceton and Hamilton Roads, in rice field, Aug. 30, 2000, DiTomaso s.n. (DAV): Edge of Middle Honcut Road, about *% mile E of Hwy 70, ca. 12 mi S of Oroville, common, valley grassland, L. Ahart 4810 (CHSC); 8 mi NE of Butte City, ditch bank. Oct. 15, 1946, J.E. Chattin and C. Ferrell s.n. (UC); Tu- lare Co.: 3 mi below Three Rivers. Sept. 25, 1920, L. Abrams 7715 (CAS): NOTES: Determination of Di- Tomaso s.n. by G.C. Tucker. The Ahart and Abrams spec- imens were originally determined as Cyperus albomargin- atus Mart. & Schrad., a synonym of C. flavicomus; how- ever, the original determinations as C. albomarginatus have not yet been confirmed. Cyperus gracilis R. Br.: DIST: SCo: CS: NCI: DOC: Los Angeles Co: volunteer in residential garden, Santa Monica. Aug. 20, 1983, Tom Yutani s.n. (CDA). Cyperus iria L.: DIST: SCo, ScV: CS: TEN: DOC: Santa Barbara Co.: Germinating in coco fiber from Sri Lanka. Commercial greenhouse in Carpenteria. Grown to maturity in CDFA greenhouse, Sacramento. Oct. 14, 1998, T. Watson s.n. (CDA); Yuba Co.: Dry to damp soil, dis- turbed area S side Woodruff Lane, just W Kimball Lane, 5 mi NE Marysville. Elev. 80 ft. Sept. 5, 1999, L. Ahart 8197 (CDA, CHSC). Cyperus owanii Boeck: DIST: CCo, SCo: CS: NCI: DOC: Los Angeles Co.: Spontaneous, UCLA Bot. Gar- den, Westwood. April 19, 1972, T.C. Fuller 1994] (CDA): San Diego Co.: Barranca, Balboa Park, San Diego. April 22, 1969, T.C. Fuller 18244 (CDA); loc. cit. Oct. 21, 1969, T.C. Fuller 19012 (CDA, DAV): San Francisco Co.: Golden Gate Park, California Academy of Sciences, San Francisco. Oct. 18, 1967, T.C. Fuller 16748; loc. cit. Nov. 21, 1968, 7.C. Fuller 18013 (CDA); Santa Barbara Co.: Smith, C.F (1976, pg. 94): NOTES: Only known occurrences in North America N of Mexico. Cyperus papyrus L.: DIST: SCo: CS: NCI: DOC: San Diego Co.: Beauchamp, R.M., (1986, pg. 48). Fimbristylis autumnalis (L.) Roem. & Schult.: DIST: ScV: CS: NW: DOC: Butte Co.: Disturbed area E side Sacramento River NW Parrott Landing, 1 mi SE Ord Fer- ry, 12 mi SW Chico. Riparian woodland. Aug. 13, 1999, L. Ahart 8145 and V. Oswald (CDA, CHSC). 94 MADRONO Scirpus cyperinus (L.) Kunth: DIST: c SNF: CS: NW: DOC: Mariposa Co.: Moist banks and alluvial flats, Mer- ced River near mouth Yosemite Creek, Yosemite Valley. Aug. 13, 1976, J.T. Howell 52233 (CAS, CDA); loc. cit. Sept. 28, 1978, G.D. Barbe 2447, 2444 (CDA, CHSC): NOTES: Determination confirmed by L. Janeway (CHSC) 2-2000. Scirpus prolifer Rottb.: DIST: SCo: CS: NCI: DOC: San Diego Co.: Persisting in sandy soil of lathhouse, 19 km NW of Fallbrook. Nov. 14, 1978, G.D. Barbe 2472 (CDA, DAV). Eriocaulaceae Eriocaulon cinereum R. Br.: DIST: SnJV: CS: EXT: DOC: Stanislaus Co.: Krause rice fields, Modesto. Sept. 18, 1947, B.G. Markos s.n. (CDA): NOTES: Extirpation likely as rice fields are no longer maintained in the Mo- desto region. Cited from California in Flora North Amer- ica (Kral 2000). Hydrocharitaceae Limnobium laevigatum (Humb. & Bonpl. ex Willd.) Heine: DIST: SCo, SnFrB: CS: NW: DOC: Alameda Co.: Jordan Pond in Garin Park; East Bay Regional Park District, Hayward. Floating and rooted in bottom mud. Osis Sig IDLO, Seo Zl, IMs, OCs ZZ, OMS Ik. Jaa s.n. (CDA); Garin Regional Park east of Hayward, silted- in pond below Jordan Pond, well-established colony both floating and stranded. Sept. 4, 1998, B. Ertter 16408 (UC); rimming Jordan pond, some in bloom, Oct. 24, 1998, B. Ertter 16458 (UC); Riverside Co.: Covering % acre of outdoor pond at Desert Lawn Cemetery, Desert Lawn Dr., Calimesa. TO2S, ROIW, Sec. 31. SB. Aug. 23, 1999, J. Chandler s.n. (CDA): NOTES: Santa Cruz Co.: in a backwater of the San Lorenzo River. Voucher specimen not yet received, but scrap material confirmed by B. Ertter in fall 2000. Not relocated in spring 2001. Floating or stranded rosettes spreading by stolons; leaves + round, 1— 3.5 cm across, sessile to long-pediceled, convexly spongy on underside. Sometimes treated as L. spongia subsp. lae- vigatum (Humb. & Bonpl. ex Willd.) Lowden. As ob- served by Ertter, eradication effort in Jordan Pond by East Bay Regional Parks District (EBRPD) has not yet suc- ceeded in eliminating the Limnobium, but has concurrent- ly severely impacted the originally diverse aquatic flora, underscoring just one of the major problems inherent in the control of aquatic pests. This plant is currently pro- hibited from being sold commercially in California; how- ever, as in the case of Salvinia molesta, it remains for sale by aquatic plant nurseries and dealers. The closely related Limnobium spongia (Bosc) Steud. (sensu stricto) is cur- rently offered for mail-order sale by midwest nurseries and is thus eventually expected to escape in California. Both taxa are a threat to California wild wetlands, irri- gation ditches, canals, sloughs, farm ponds or private lakes, and if either should become established in navigable waterways they are especially likely to spread rapidly and widely. The two established and reproducing colonies de- scribed above are in non-navigable waters. It should also be noted that any number of aquatic taxa sold unlabeled in California nurseries could be Limnobium or other un- identified taxa which could display the same degree of aggressiveness as does Limnobium should they be intro- duced into aquatic habitats where there are no natural con- trols. [Vol. 49 Iridaceae Tris foetidissima L.: DIST: NCoRO, SnFrB: CS: N: DOC: Alameda Co.: well established along a stretch of the Golden Spike Trail in Redwood Regional Park, Oak- land Hills. Oct. 15, 2000, Ertter 17542 (UC); Contra Costa Co.: Sibley Volcanic Preserve in Oakland Hills, single clump in understory of oak-pine forest behind in- terpretive center. May 20, 2000, B. Ertter 17025 (UC); Sonoma Co.: Best, C., et al. (1996, pg. 261): NOTES: Spreads readily in garden situations. Seeds with red arils indicate a high potential for bird dispersal into wildlands. Tris germanica L.: DIST: NCoRO, SnFrB: CS: TEN: DOC: Santa Barbara Co.: Munz, P.A. (1974, pg. 907); Sonoma Co.: Best, C., et al. (1996, pg. 261): NOTES: Seen often about old habitations and sometimes persistent long after signs of human infestation have disappeared. Citations here both imply local naturalization. Also ob- served as a casual in Alameda and Contra Costa Cos. Tris orientalis L.: DIST: SCo: CS: NCI: DOC: Santa Barbara Co.: Smith, C.F (1976, pg. 106). Ixia polystachya L.: DIST: NCo: CS: NCI: DOC: Mendocino Co.: persisting in scattered locations among the headstones of Evergreen Cemetery, Hwy | at Moun- tain View Rd, | mi S of Manchester. T13N, R17W, Sec. 06, MD. May 27, 1982, G.D. Barbe 3628 (CAS, CDA): NOTHES: dupl. det. by J.-M. Mullin, 1983 (BM). Ixia speciosa Andrews: DIST: NCo: CS: NCI: DOC: Mendocino Co.: Smith, G. and C. Wheeler (1990-1991, Pearls): Moraea collina Thunb.: DIST: CCo: CS: N: DOC: Santa Cruz Co.: remnant native coastal prairie site on marine terrace lying between Rodeo Gulch and Hidden Valley Road, ca. 75 meters SSE of pole 14/117 of the PG and E Paul Sweet-Green Valley 115 kv double-wood pole transmission line (site is 1.5 air mi NNE of Dominican Hospital); elev. 370 ft, grassland on edge of thickets of Quercus agrifolia. May 2, 2000, D.W. Taylor 17446 (JEPS): NOTES: Naturalization localized at present. Goldblatt (1998) treats this as Moraea collina Thunb., not as Homeria (Thunb.) Salisb. which is the name in use by the horticultural trade and by Australian authors. The en- tire genus Homeria is on the Federal Noxious Weed Act quarantine list (see Federal Register, May 25, 2000, p. 33741-33743), and is thus illegally sold, although com- monly available in California nurseries. Native to South Africa. Related species are aggressive invaders of pastures in New Zealand, Australia. All Homeria spp. are toxic to livestock. Moraea polystachya Ker Gawl.: DIST: SCo: CS: NCI: DOC: Santa Barbara Co.: Plants abundantly naturalized on roadside, N side of Cabrillo Blvd., SW corner of Bird Lagoon, Montecito. TO4N, R27W, Sec. 24, SB. Dec. 11, 1968, 7.C. Fuller 18041; loc. cit. Mar. 23, 1971, T.C. Fuller 19820 (CDA). Juncaceae Juncus nodatus Cov.: DIST: SnFrB: CS: N: DOC: Alameda Co.: UC-Berkeley Botanical Garden, edge of path near vernal pool section. July 16, 1998, H. Forbes s.n. (UC): NOTES: Well-established local colony, not de- rived from cultivated material. Determination by B. Ertter. Liliaceae (sensu lato) Agapanthus praecox Willd.: DIST: CCo: CS: NCI: DOC: San Mateo Co.: McClintock, E., et al. (1990, pg. 167): NOTES: Clearly originating as a garden escape. 2002] Allium cepa L.: DIST: NCoRO: CS: NCI: DOC: Son- oma Co.: Best, C., et al. (1996, pg. 269): NOTES: A local garden escape in 1963, probably not persistent. Allium sativum L.: DIST: NCoR: CS: C: DOC: Son- oma Co.: Best, C., et al. (1996, pg. 269): NOTES: A local garden escape. Amaryllis belladonna L.: DIST: CCo, SCo, SnFrB, SnBr: CS: TEN: DOC: Contra Costa Co.: observed by Ertter near Donner Cabin, Mount Diablo State Park [voucher prepared but lost]; observed elsewhere in San Francisco Bay Area.; Los Angeles Co.: Malibu, upslope from Winter Canyon, near Pepperdine Univ., N of Pacific Coast Hwy, E of Malibu Cyn Rd., SW of Malibu Civic Center Way, elev. ca. 300 ft, 34°02’N; 118°42’W, TOIS, R17W, Sec. 31, SB. Mar. 15, 1995, S. White 2585 (UCR); Monterey Co.: Yadon, V. (1995); San Mateo Co.: McClintock, E., et al. (1990, pg. 167); Sonoma Co.: Sea Ranch, just ESE of Leeward Rd. entrance along State Hwy. 1. Aug. 13, 1975, M. Hektner 220 (DAV), voucher for Wasmann J. Biol. 35(1):26—-53: NOTES: Also ob- served by Sanders apparently naturalized near Yucaipa, San Bernardino Co. Cultivated widely. Often long-per- sistent from cultivation and known from numerous home- stead sites in northern and southern California. Chlorophytum capense (L.) Druce: DIST: CCo: CS: C: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p-53); Thomas, H. (1961, p. 117): NOTES: A garden es- cape. Hyacinthus orientalis L.: DIST: CCo: CS: C: DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 53); Thomas, H. (1961, p. 117): NOTES: A garden escape. Kniphofia uvaria (L.) Hooker [or hybrid derivitive]: DIST: CCo, NCo, SCo, SnFrB: CS: NW: DOC: Alameda Co.: North foot of Albany Hill (eliminated by subsequent roadwork). May 25, 1992, B. Ertter 11070 (UC); Contra Costa Co.: Miller Knox Regional Park, open hillside south of watertank, single large patch, expanding. Mar. 21, 1999, B. Ertter 16487 (UC); Humboldt Co: Arcata Marsh and Wildlife Sanctuary, dike at Klopp Lake, on disturbed fill on Humboldt Bay. April 9, 2000, G. Leppig 1287 (CDA, HSC); Mendocino Co.: Smith, G. and C. Wheeler (1990-1991, pg. 107); Hwy. 1 at Navarro Rd, 7.5 miles S of Elk. Roadside pastures. Common. July 9, 2000, G. Leppig 1459 (CDA, HSC); Sinkyone Wilderness State Park, mouth of Jackass Creek at the end of Wheeler Rd. Old home site on the S side of Cyn. T23N, R19W, Sec. 01, MD. May 28, 1987, F. Bowcutt 1011 (DAV); San Francisco Co.: Howell, J.T. et al. (1958, pg. 53); San Mateo Co.: McClintock, E., et al. (1990, pg. 168);. Santa Barbara Co.: upper east fork of Cold Spring Canyon above Montecito. Jan. 21, 1967, H.M. Pollard s.n. (SBBG, UCR); Sonoma Co.: Best, C., et al. (1996, pg. 275): NOTES: Expected elsewhere. Can be long persistent from cultivation as at Fort Hunter Liggett (Monterey Co.) where it is found about old homesteads without any in- dication of spontanaety. Other ‘populations’ may originate via garden trash, but many locations are obviously spon- taneous. Observed in coastal meadows and on roadsides, where occasional to common in Monterey Co. but not yet vouchered. Also observed in locations other than the records above in Sonoma, Mendocino and Humboldt Cos. Leucojum aestivum L.: DIST: CCo: CS: NCI: DOC: San Luis Obispo Co.: Keil, D.J., et al. (1985, pg. 222). Narcissus pseudonarcissus L.: DIST: NCo, SnFrB: CS: NW: DOC: Contra Costa Co.: Mount Diablo State Park, Donner Cabin site, persisting colony. Feb. 16, 1997, B. Ertter 15451] (JEPS); northwest end of Wildcat Canyon Park, occasional clumps among brush-grassland mosaic. HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 95 Mar. 7, 1999, B. Ertter and L. Fujii 16485 (UC); Molate Beach Park, coastal prairie. April 7, 1996, B. Ertter 1458] (UC); Humboldt Co.: Fickle Hill Road, abundant in open areas. TOON, ROIE, Sec. 27, H. Mar. 9, 1963, J.C. Reppas 18 (HSC); Old Arcata Road at Jacoby Creek Rd., common along roadsides, TOSN, ROIE, Sec. 16, H. Apr. 8, 1965, N.D. Dennis 15 (HSC); North Bank Rd., 4 mi N of Arcata, escaped from cultivation, ROIE, TOON, Sec. 09, H. Apr. 14, 1965, J. L. Baker 1965 (HSC): NOTES: A diversity of hybrid cultivars are also sporadically encountered (e.g., Molate Beach Park, Contra Costa Co.) These may be persistent from cultivation, the remnants of garden trash, or spontaneous. Narcissus tazetta L.: DIST: NCo, SnFrB: CS: NW: DOC: Contra Costa Co.: Mount Diablo State Park, Back Canyon Road at cross road above Donner Canyon trail- head, seepage area in grassland, well established spread- ing colony. Jan. 25, 1998, B. Ertter and A. Dennis 15910 (JEPS); Humboldt Co.: Patrick’s Point State Park, near Ceremonial Rock. 21 July 1986. E. Mackey s.n. (HSC). Ornithogalum umbellatum L.: DIST: CCo, NCoRI: CS: GH/C: DOC: Lake Co.: Upper Lake, along Menden- hall Rd, off Hwy 20, in walnut orchard. Apr. 4, 2000, R. Elkins s.n. (DAV); Santa Cruz Co.: Garden weed, Riv- erside Dr., Watsonville. Apr. 12, 1971, D.H. Shaw s.n. (CDA). Pancratium maritimum L.: DIST: SCo: CS: N: DOC: Los Angeles Co.: El] Segundo Dunes, immediately west of L.A. International Airport and Pershing Dr., E of Vista Del Mar Blvd., nr. 33°56'N; 118°26'W, coastal dunes for- merly largely occupied by residential neighborhoods, the houses removed ca. 15 years ago. Assoc. with Croton cal- ifornicus, Abronia umbellata, Camissonia cheiranthifolia and various persisting ornamentals. Forming vegetative clumps in sand; also some reproduction by seed. July 22, 1987, A.C. Sanders 7156 (RSA, SBBG, UCR): NOTES: An escaping ornamental; native to beaches in southern Europe. Reported also from Ventura Co. at Mugu Lagoon and San Buenaventura State Beach, but no specimens yet available for confirmation. Tulipa clusiana DC. in Redoute: DIST: SCo: CS: TEN: DOC: Riverside Co.: City of Riverside, east edge near Moreno Valley, spontaneous at the site of the former Desert Nursery, N of Hwy 60, E of Day Street, unknown origin, possibly accidentally introduced with other plant material, now reproducing and increasing in one area with no care or watering. Mar. 23, 1994, A. Miller 94-1 (UCR): NOTES: At almost the same time as this collection, this species was reported to Sanders as a lawn weed in Riv- erside, but no voucher materialized. This plant is appar- ently a common weed in the Mediterranean basin. It is quite showy and might get moved around intentionally as an ornamental. Poaceae Acrachne racemosa (Roem. & Schult.) Ohwi: DIST: SCo: CS: TEN: DOC: Riverside Co.: Sanders, A.C., (1996, pg. 524). Aira caryophyllea L. var. cupaniana (Guss.) Fiori: DIST: SnFrB: CS: NCI: DOC: Contra Costa Co.: Mount Diablo State Park, Wall Point Road, prescribed burn. May 28, 1995, M.L. Bowerman s.n. (JEPS), verified by J. Wipff, 1998: NOTES: =Aira cupaniana Guss. Not relo- cated in 1998—99. Amphibromus neesii Steud.: DIST: ScV: CS: NW: DOC: Sacramento Co.: grown to maturity in greenhouse from plant dug out of large colony in vernal pool at corner 96 MADRONO of Sunrise and Keifer Aves., SE of Sacramento. July 28, 2000. G.F. Hrusa s.n. (CDA): NOTES: Although known from only a single site, this species is of particular interest because of its ability to invade and survive in vernal pools. Native to and sometimes listed in the Australian literature under the synonym Amphibromus nervosus (R. Br.) Druce. First seen in this site in 1990 by J. Glazner of North Fork Associates, Auburn, CA (personal communi- cation to Hrusa). Amphibromus neesii forms cleistoga- mous seeds; these first noted and identified at the Cali- fornia Dept of Food and Agriculture Analysis and Iden- tification (now Plant Pest Diagnostics) Branch Seed Lab- oratory in 1967 as a contaminant in Trifolium subterraneum seed imported from Australia. Aristida dichotoma Michx.: DIST: n SNF: CS: NW: DOC: Butte Co.: Dry bare granite soil, Big Bald Rock. Uncommon, inconspicuous. Yellow pine forest, elev. 3260 ft. Aug. 5, 2000, L. Ahart 8623 and V. Oswald (CDA, CHSC); Dry bare granite soil above seeps on bare granite. N side Bean Ck. Rd., ~ %4 mi. SE Little Bald Rock and 1% mile NE Big Bald Rock, 4 airmiles NE Madrone Lake and 13 mi NE of Oroville. Uncommon. Yellow pine for- est, elev. 3500 ft. T21N RO6E, Sec. 29, MD. Aug. 5, 2000, L. Ahart 8624 and V. Oswald (CDA, CHSC): NOTES: Determinations by K. Allred (NMCR). Chloris truncata R. Br.: DIST: DSon, SCo, SnJV: CS: N: DOC: Imperial Co.: Weed in alfalfa, 5 mi S of El Centro and 3.5 mi. SW of Heber. T16S R13E, Secs. 35, 36. Aug. 8, 2000, J. Johnson s.n. (CDA, DAV); Merced Co.: Almond orchard at corner of Vista Ave and Hwy. 99 S, N of Chowchilla. July 19, 1999, J. DiTomaso s.n. (CDA, DAV); Riverside Co.: Sanders, A.C., (1996, pg. 526; 1999, pg. 113); Weedy in hybrid bermudagrass turf, turfgrass nursery, Leon Rd. Winchester. Nov. 9, 1978, E. Storm s.n. (CDA), det. by G.E Hrusa, Oct., 1999. Echinochloa crusgalli subsp. spiralis (Vasing.) Tzvel- ev: DIST: ScV, SnGB: CS: N: DOC: Butte Co.: Afton Rd. nr. Biggs. Aug. 23, 1947, M.K. Bellue s.n. (CDA); In milo field. Sept. 16, 1958, J. Harroun s.n. (CDA); Los Angeles Co.: San Gabriel Mtns., Angeles National Forest: Little Rock Reservoir, dry sunny sandy flat toward upper end. Sept. 6, 1966, L.C. Wheeler 8873 (CDA, RSA); Sac- ramento Co. sine loc. July, 1943, Bellue s.n. (CDA): NOTES: First report of this variety for North America. Expected widely. Determinations by S.L. Mosyakin (KW), March 26, 2001. Echinochloa esculenta (A. Br.) H. Scholz: DIST: NCo, SCoRI, SnJV: CS: NCI: DOC: Del Norte Co.: Abundant along a slough, Reservation Ranch, Smith Riv- er. Oct. 2, 1968, E.J. Garrett s.n. (CDA); Monterey Co.: near Chualar. Sept. 1938, McElrath s.n. (CDA); Fresno Co.: near Fresno. Sept., 1938, McElrath s.n. (CDA): NOTES: Determinations by S.L. Mosyakin (KW), March 26, 2001. Echinochloa frumentacea Link (E. crusgalli var. frumentaceum (Link) Trimen) has been consistently mis- applied to this plant in California. It is probably most often an escape from cultivation, although populations may be locally persistent. Ehrharta longiflora Sm.: DIST: SCo: CS: NW: DOC: San Diego Co.: Simpson, M.G., et al. (1996, pg. 79): NOTES: Known only from a single location. Eragrostis curvula (Schrad.) Nees var. conferta Nees: DIST: SCo: CS: N: DOC: Riverside Co.: Sanders, A.C. CIGRNO. (2s DZ). Eremochloa ciliaris (L.) Merr.: DIST: CCo: CS: EXT: DOC: San Francisco Co.: Thurber, G. in S. Watson, (1880, pg. 261—2): NOTES: Original report as Jschaemum leersioides Munro (specimen at CAS). Citation in Howell, [Vol. 49 J.T. et al. (1958), is based on the above report. Extirpation is assumed although there has been no thorough investi- gation of its presence or absence. Gaudinia fragilis (L.) P. Beauv.: DIST: NCoRO: CS: NW: DOC: Sonoma Co.: Daniel, T. and C. Best (1992, pp. 309-310); Best, C., et al. (1996, pg. 300): NOTES: Reported location highly localized, should be sought else- where in the region. Glyceria fluitans (L.) R. Br.: DIST: CaRH, NCo, NCoRI: CS: NW: DOC: Del Norte Co.: Veneer Mill pond, Redwood NP. June 5, 1995, G. Leppig 242 (CDA, HSC); Humboldt Co.: Arcata Bottom, wet slough. July 20, 1933, J. T. Tracy 12801 (HSC, RSA); Stone Lagoon moist field. Aug. 3, 1924, J. T. Tracy 6749 (RSA); Arcata, Alder Grove Pond, on edge of pond. June 4, 1995, G. Leppig 242 (CDA, HSC); loc. cit. May 25, 1998, G. and S. Leppig and K. Neander 791 (CDA, HSC); loc. cit. April 24, 1999, G. and S. Leppig 1125 (CDA, HSC); Lake Co.: Kelseyville, swampy places. June 8, 1924, J.W. Blankin- ship s.n. (RSA); Shasta Co.: Cassel. June 24, 1930, M. Kjilsberg s.n. (DAV): NOTES: According to G. Leppig (HSC), this may be an overlooked native. Hordeum vulgare L. [sensu lato. Including material re- ferable to var. trifurcatum (Schlecht.) Alef.]: DIST: CA- FP: CS: C: DOC: Alpine Co.: W Fk. Carson River, ca. 1 mi W from Woodfords along Crystal Springs Rd. June 22, 1974, D.W. Taylor 3911 (UC); Los Angeles Co.: San Cle- mente Island, widely scattered over mesa summit and S slopes. May, 1936, N. Murbarger 174 (UC); Marin Co.: nr. Inverness. June 3, 1945, J.T. Howell 20918 (UC); Mariposa Co.: valley floor, Yosemite Valley, Yosemite Natl. Park. Apr. 1, 1934, P.S. Bartholomew s.n. (UC); Mendocino Co.: nr. Ukiah, volunteer in oat-field. May 25, 1899, J. Burtt Davy and W.C. Blasdale 5059 [awned] (UC); May 25, 1899, J. Burtt Davy 5058 [awned] (UC); Sacramento Co.: Volunteering in residental garden. Mar. 29, 1979, K. Miller s.n. (CDA); San Diego Co.: Simpson, M.G. et al. (1996, pg. 80); San Luis Obispo Co.: ROW along Hwy 46 approx. 4 mi E Paso Robles. Weed on roadside, solitary. Elev. 200 m. 35°39’N; 120°26’W. Apr. 4, 1992, G.F. Hrusa 10345 (CDA); Cuyama Valley, weed at edge of barley field, immed. W of int. Hwy 33 and Hwy 166. Penetrating into planted field along edges. Elev. 650 m. 34°55'N; 119°33’W. Apr. 26, 1998, G.F. Hrusa 14276 (CDA); Siskiyou Co.: Volunteer; subsaline flat and roadside, betw. Old Highway 99 and RR at Truttman Lane, 3.2 km S of Grenada. Elev. 800m. T44N, RO6W, Sec. 34, MD. Apr. 28, 1980, G.D. Barbe 2589 (CDA); Ventura Co.: San Nicolas Island, N edge of mesa, E end of old landfull, E of Living Compound. Apparently used for erosion control on inactive landfill site. Apr. 1, 1992, S. Junak SN-789 (JEPS, SBBG): NOTES: Although spo- radic occurrences are well-documented, these were all de- termined to be casuals and accordingly the species was not included in The Jepson Manual. This species is also commonly used for roadcut erosion control and it may be difficult to determine if a local site or population is estab- lished or recently planted. Records listed above are rep- resentative; the species can be expected in every Califor- nia county. Numerous unvouchered records are on file at CDA (based on specimens submitted but not retained); observed as common on road cutbanks throughout north- ern California, especially so along that part of the Inter- state 5 corridor. Hordeum vulgare var. trifurcatum is a hooded form, reported as a roadside weed almost as often as the typical variety, although probably due to its visible distinctiveness and not frequency. Annotations at UC by N. Jacobsen (1980) do not recognize var. trifurcatum. 2002] Leptochloa dubia (Kunth) Nees: DIST: CaRF/n SNF: CS: N: DOC: Butte Co.: west side of Hwy 70 ca. 1% miles north of bridge across the West Branch of Lake Oroville, just north Rich Gulch Road, T22N, RO4E, Sec 09, elev. 1300 ft. Common on dry rocky disturbed soil. Sept. 6, 1997, L. Ahart 7894 (CHSC, JEPS). Nassella tenuissima (Trin.) Barkworth: DIST: CCo: CS: C: DOC: Contra Costa Co.: Creekside Park, El Cer- rito, overflow basin for Cerrito Creek, single vigorous plant. July, 2000, B. Ertter s.n. (UC): NOTES: Also read- ily establishing in sidewalk cracks adjacent to cultivated plants in gardens throughout Berkeley, this species has a high potential for spread into natural areas. A popular hor- ticultural species whose weedy behavior should be care- fully monitored. Panicum maximum Jacq.: DIST: SCo: CS: GH/C: DOC: Riverside Co.: Sanders, A.C. (1996, pg. 529). Panicum repens L.: DIST: SnFrB: CS: NCI: DOC: Alameda Co.: abundant weed of ornamental shrubs, climbing to 3 ft high in some shrubs, Oxford Tract, U.C. campus, NE corner of Walnut St. and Hears Ave., Berke- ley. TOIS, RO4W, Sec. 02, MD. Nov. 21, 1968, T.C. Ful- ler 18011 (CDA): NOTES: Rhizomatous perennial intro- duced into California from Florida in Melaleuca stock. Panicum rigidulum Bosc ex Nees var. rigidulum: DIST: ScV, n SnJV: CS: NCI: DOC: Butte Co.: Bank of Feather River, Oroville; elev. 175 it. Sept. 22, 1961, J.T. Howell 36714 (CAS, CDA), original det. as P. agrostoides Spreng.; Sacramento Co.: 1 mi S of Fair Oaks along American River. Growing in sandy gravel of dry flood banks along river. Oct. 31, 1961, B. Crampton 6072 GabiW@): Moc: cit: Oct, 116; 1953.4 B Crampton 1706 (AHUC); American River near Natoma. Moist sandbar. Sept. 21, 1950, R. Tofsrud s.n. (AHUC); Stanislaus Co.: edge of Tuolumne River 3 miles SW of La Grange, wet soil. Aug. 23, 1961, P. Allen s.n. (JEPS); Tuolumne Co.: W side of Hetch Hetchy Dam, Swamp Lake Area, Yosem- ite NP. Aug. 9, 1958, H.L. Mason 14855 (UC); Yolo Co.: Common among rocks, east levee of Merritt Island, at Rd 142, SE corner of County. July 22, 1972, C. Quick 72-12 (AHUC): NOTES: Current determination of P. Allen s.n. by M. G. LeLong, 1995; previous determinations include P. lindheimeri Nash. and P. agrostoides Spreng. Included in Oswald and Ahart (1995, pg. 290) as a synonym of P. acuminatum var. lindheimeri (Nash) Fern. Panicum texanum Buckl.: DIST: SnJV: CS: TEN: DOC: Fresno Co.: In a vineyard along Academy Ave. near Sanger. August, 1983, B. Fischer s.n. (AHUC), det. by B. Crampton, 1983; Kern Co.: Weed in vineyard, De- lano. Sept., 1992, H. Kempen s.n. (DAV), det. by G.FE Hrusa, 1992: NOTES: An uncommon agricultural weed, sometimes treated in Urochloa, and so filed at DAV. Pennisetum glaucum (L.) R. Br.: DIST: SnJV: CS: C: DOC: Merced County: Highway 33 about 3 miles S of southern outskirts of Gustine; moist roadside ditch near Main Canal at Pfitzer Road; TO8S, RO9E, Sec 29, MD. 37°12'28"N; 121°00'37"W; Howard Ranch quad., elev. 110 ft. With Paspalum dilatatum, Sorghum halepense dominant. July 12, 2000, D.W. Taylor 17480 (JEPS): NOTES: Escape from cultivation? Large areas were being grown (presumably for dairy silage) within about a half mile upstream of the site above. Although there was a sizable population, the occurrence could be attributed to seed washing downslope along the ditch alignment. In the Jepson Manual (pg. 1296), this name was misapplied to Setaria pumila (Poir.) R. & S. as the synonymized com- bination Setaria glauca (L.) P. Beauv. Pennisetum latifolium Spreng.: DIST: CCo: CS: NCI: HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 97 DOC: Santa Cruz Co.: three clumps of plants to 12 ft, naturalized on shoulder of Southern Pacific RR just N of 30th Ave. crossing, Santa Cruz. Sept. 29, 1969, J. Bauer si =(CDA); oc: cit. ‘Oct. 19, 1969, T.C. £uller’ 18980 (CDA, DAV, UC); Yolo Co.: Davis, UC farm, volunteer in row A12 of grass garden. Oct. 25, 1923, P.B. Kennedy s.n. (AHUC). Phalaris coerulescens Desf.: DIST: SnFrB: CS: C: DOC: Contra Costa Co.: Mount Diablo, Black Hawk Ridge Road south of Sycamore Creek, single plant on roadside. June 12, 1999, B. Ertter 16713 (JEPS): NOTES: Like P. paradoxa, but perennial with bulbous based culms, sterile spikelets not clublike. Not relocated in 2000, possibly due to road grading at the site. Phyllostachys aurea A. & C. Riviere: DIST: SCo: CS: NCI: DOC: Placer Co.: Open dry slope above wash, 2 mi W of Auburn. Nov. 13, 1969, B. Crampton 8531 (AHUC); Stanislaus Co.: Colony established along per- manent stream, La Grange Dam Rd | mi E of La Grange. June 20, 1969, P.S. Allen 429 (DAV); Ventura Co.: Pop- lar grove and thicket at juncture of San Antonio Creek and Ventura River near Arnaz Apple Orchards, Oak View. Re-collected after 15 yr in same spot. Colony enlarged. Dec. 21, 1962, H.M. Pollard s.n. (AHUC, CAS, CDA, SBBG). Phyllostachys bambusoides Siebold & Zuccarini: DIST: CCo: CS: NCI: DOC: San Mateo Co.: Mc- Clintock, E., et al. (1990, pp. 181-182). Piptochaetium stipoides Hackel ex Arech. sensu lato: DIST: NCo: CS: NW: DOC: Marin Co.: About 4 mile E of Hwy 1 on trail to Bolinas Ridge, just north of road to Alpine Dam. May 9, 1978, C. Best s.n. (CAS, CDA); single plants or patch to 3 meters across, S-facing slope of canyon on W slope of Bolinas Ridge, 0.5 km E of Hwy 1 at southern limits of Dogtown. TOIN, RO8W, Sec. 11, MD. June 2, 1978, G.D. Barbe 2392 (CDA); loc. cit. G.D. Barbe 2391 (cited in correspondence, specimen location unknown), determinations by Gladys Perez-Camargo (BAA), B. Rosengurtt (MVFA), and L.T. Ellis (K); infre- quent patches to 10 meters across, S facing slope W of Bolinas Ridge, betw. Coppermine and Wilkins gulches at Dogtown (Woodville), 4 km N of Bolinas. Elev. 121 m. TOIN, RO8W, Sec. 11, MD. May 16, 1979, G.D. Barbe 2502 (CDA, DAV, UC); Ridge between Garden Canyon and Pike County Gulch on western slope of Bolinas Ridge ca. 3.25 miles north of Stinson Beach. Occasional in hard packed soil of old road. May 17, 1978, 7.H. Harris s.n. (DAV): NOTES: Numerous more recent collections from the same localities above are at UTC. It is interesting that although the original California specimens of the P. sti- poides alliance were determined as var. purpurascens (Hackel) Parodi, study of the associated correspondence indicated considerable variation among these specimens. Intraspecific application within the P. stipoides group is currently in dispute; and in addition, Barbe 2392 was orig- inally det. at K as P. grisebachii (Speg.) Herter. Subse- quent annotations at SI by E.G. Nicora indicate all were part of P. stipoides var. purpurascens. More recent col- lections determined by E. Sanchez (BA) again indicate two taxa are present at this site: P. stipoides var. purpur- ascens and var. stipoides. Apparently further field study on Bolinas Ridge would be useful. Well-established there where it is mixed with P. setosum. The land was private with restricted access until two years before the first col- lections in 1978, thus the species has possibly been pre- sent on the site for a considerable time. Speculated to have been introduced with cattle from S. America. Polypogon imberbis (Phil.) Bjorkm.: DIST: CCo: CS: 98 MADRONO NCI: DOC: Contra Costa Co.: near Martines. June 7, 1900, Burtt Davy 6662 (US); San Luis Obispo Co.: Oceano Beach in moist spots among sand dunes. July 18, 1947, R.F. Hoover 7314 (OBI, US): NOTES: Included in the Jepson Manual as P. elongatus Kunth, based on the two specimens cited above. Both of these recently re-de- termined by Robert Soreng (US) as P. imberbis. Hoover for San Luis Obispo Co. (1970) and especially Mason (1957) were uncomfortable with the P. elongatus deter- minations but did not suggest an alternative name. Pseudosasa japonica (Sieb. & Zucc. ex Steud.) Ma- kino ex Naka: DIST: CCo: CS: NCI: DOC: San Fran- cisco Co.: Howell, J.T. et al. (1958, pg. 43); Thomas, H. (1961, p. 75): NOTES: Spreading vegetatively from cul- tivation. Schedonnardus paniculatus (Nutt.) Trel.: DIST: CaR: CS: NCI: DOC: Siskiyou Co.: fenceline grass, Ager Road, Montague. Aug. 5, 1980, D.H. Shaw s.n. (AHUC, CDA). Spartina anglica C.K. Hubb.: DIST: CCo: CS: NW: DOC: Marin Co.: Creekside Park adjacent to Corte Ma- dera Creek, Greenbrae. Growing in low-flow channel of coastal salt marsh, assoc. with Distichlis spicata and Spar- tina densiflora. May 6, 2000, S. Klohr s.n. (CDA, DAY): NOTES: Only known site in California. More likely in- troduced to Creekside Park during marsh restoration work in 1977 sensu Spicher and Josselyn (1985) than via natural dispersal from Washington sensu Bossard et al. (2000). Stipa capensis Thunb.: DIST: DSon (Coachella Val- ley): CS: NW: DOC: Riverside Co.: Coachella Valley/ San Jacinto Mtns. Foothills. Alluvial fan of Chino Canyon along the Palm Springs tram road, N of the road 14 tele- phone poles above Hwy 111 (Palm Canyon Dr.), elev. 300 m/950 ft, 33°51’N; 116°34'W, TO4S RO4E Sec. 04, SB. Creosote bush scrub on rocky alluvium cut by arroyos; with Larrea, Hymenoclea, Hyptis, Ambrosia, Justicia, etc., three patches seen, ca. 200—300 individuals. Mar. 19, 1995, A.C. Sanders 16148, with G. Helmkamp, P. MacKay, et al. (UCR), det. by M. Barkworth; loc. cit. Apr. 6, 1995, A. C. Sanders and M. Skinner 16393 (UCR); Coachella Valley, Chino Cyn., foothills of the San Jacinto Mtns., along the road to the Palm Springs Tram 1.5 mi above Hwy. 111. Palm Springs 7.5’ quad., 33°50'34"N; 116°34'51”"W, TO04S, RO4E, Sec. 04, SB. Elev. 1200 ft/366 m, rocky loam on alluvial fan, creosote bush scrub with Larrea, Ambrosia dumosa, Opuntia echinocarpa, Kra- meria grayi, etc., common annual on roadside and spread- ing into desert vegetation. Apr. 15, 2000, A.C. Sanders 23321, with Giles Waines, Mitch Provance, T.B. Salvato, et al. (UCR); Cathedral Canyon, border of Rancho Mirage [and] Cathedral City, 33°45’N; 116°30’W. Mar. 11, 1997, Denise Woodard and Gilbert Goodlet s.n. (UCR), det. by A.C. Sanders; San Jacinto Mtns., S of Chino Canyon, at NW end of Palm Springs, 33°50'20"N; 116°33'45"W, TO4S, RO4E, Secs. 09 and 10, SB. Elev. 1148 ft/350 m, flat areas with Hyptis, Psorothamnus schottii, some creo- sote bush and smoke trees, locally abundant in disturbed places. Mar. 18, 1997, J. Wear and N. Moorhatch s.n. (UCR), det. by A.C. Sanders: NOTES: First records for California and North America of this annual Stipa with long awns and sharp callus tips. This species will be a severe nuisance if it becomes widely established. The seeds readily become caught in the fur of dogs and other animals and so will probably create veterinary problems, and will certainly be subject to ready dispersal. The very sharp callus can easily pierce human skin and cause un- pleasant sores. This plant is obviously a Stipa in the broad [Vol. 49 sense, but its distinctly annual habit will quickly distin- guish it from all other known Californian Stipeae. Acc. to M. Barkworth, in the narrow taxonomic sense this plant is an Achnatherum, but the published combination in that genus by P. Beauv (Essai Agrostogr. 146) is invalid, hav- ing as its basionym Milium capense L. and not Stipa ca- pensis Thunb. Thus, in Achnatherum there is not currently an available epithet. Themeda quadrivalvis (L.) Kuntze: DIST: SCo: CS: TEN: DOC: San Bernardino Co.: City of Ontario; pri- vate residence, southeastern corner of yard beneath power lines. Nov. 24, 1991, T7.S. Ross 6026 (CDA, RSA). Tribolium obliterum (Hemzl.) Renvoize: DIST: CCo: CS: NW: DOC: Monterey Co.: well-established on sand dunes in scattered locations about the former Fort Ord Army base. June 6, 2000, B. Delgado, E. Finley, B. Oliver s.n. (CDA, DAV); Matthews, M.A. (1997, p. 341); loc. cit. April 17, 1990, V. Yadon H-3828 (PGM); loc. cit. April 14, 1998, V. Yadon H-3695 (PGM): NOTES: First collected by V. Yadon (PGM). Sieglingia decumbens (L.) Bernh. [=Danthonia decumbens (L.) DC.] misapplied. Reported in Matthews (1997) under that name. First re- cords for North America. Native to South Africa. Triticum aestivum L.: DIST: CA-FP: CS: C: DOC: Alameda Co.: Anthony Chabot Regional Park, archery range. June 2, 1981, J. Stratford s.n. (JEPS); Fresno Co.: Big Creek, 5000 ft. Aug. 9, 1956, H.M. Pollard s.n. (DAV, SBBG); Humboldt Co.: Arcata waste area near Com- munity center. Apr. 14, 2000, G. Leppig 1293 (CDA, HSC); Kern Co.: Twisselmann (1956, pg. 211); Los An- geles Co.: San Gabriel Mtns, Glendora Mtn. Rd., int. with Upper Monroe Rd. Rd. margin and adjacent vegetation. Locally naturalized. May 8, 1992, T. Ross 6305 (UC); Marin Co.: Penalosa, J., (1963, pg. 27); Howell, J.T. (1970, p. 74); Mendocino Co.: nr. Walkers Valley. May 25, 1899, J. Burtt Davy and W.C. Blasdale 5062 (UC); Monterey Co.: roadside on Dolon Rd at Elkhorn Rd, May 24, 2000, G. Leppig 1383 (CDA, HSC); San Francisco Co.: Howell, J.T. et al. (1958, pg. 45); San Luis Obispo Co.: Appearing from straw used to stabilize planted areas on sand dunes immed. W of Oso Flaco Lake. Elev. 150 ft. Apr. 5, 1987, A.P. Griffiths 18086 (CDA, OBI); loc. cit. May 18, 1987, A.P. Griffiths 18187 (CDA, OBI); San Mateo Co.: McClintock, E., et al. (1990, pg. 184); Santa Barbara Co.: Santa Barbara, N of Botanic Garden, Mis- sion Cyn. Cr. June 19, 1944, C.F. Smith 945 (DAV, SBBG); Sonoma Co.: Best, C., et al. (1996, pg. 312): NOTES: The above list is not exhaustive. As a casual, this species is relatively commonly encountered and can be expected throughout California; it is likely known from every county. Probably the most common introduction vector is straw bales used for roadside erosion control, but also may be a contaminant or a component of seeding mixes. It is popular for use as the latter because of its usually ephemeral nature. Although individual sites rarely persist, new locations appear regularly. Naturalization has been reported however, as in Ross 6305 above (Los An- geles Co.), and its extent should be investigated further. Pontederiaceae Heteranthera rotundifolia (Kunth) Griseb.: DIST: ScV: CS: N: DOC: Butte Co.: Oswald, V.H. and L. Ahart, (1994, pg. 297); Yuba Co.: Damp soil in rice field, S of Woodruff Lane, just E of Mathews Rd, E of Sam Shin- taffer’s rice dryer, approx. 5 mi NE of Marysville. Com- mon. Elev. 80’. TI6N, RO4E, MD. Sept. 17, 1999, L. Ahart 8241 (CDA, CHSC). MAprRONO, Vol. 49, No. 2, pp. 99-114, 2002 FIELD ASSESSMENT OF THE CALIFORNIA GAP ANALYSIS PROGRAM GIS DATABASE IN CENTRAL CALIFORNIA JOHN E KARLIK' EUGENE D. ALBERTSON University of California Cooperative Extension, 1031 S. Mt. Vernon, Bakersfield, CA 93307 Y. JAE CHUNG U.S. Army Corps of Engineers, Los Angeles District, Regulatory Branch, PO. Box 532711, Los Angeles, CA 90053-2325 ALISTAIR H. MCKAY University of California Cooperative Extension, 1031 S. Mt. Vernon, Bakersfield, CA 93307 ARTHUR M. WINER Environmental Science and Engineering Program, 650 Young Drive, University of California, Los Angeles, CA 90095-1772 ABSTRACT Given the key role played by biogenic volatile organic compounds (BVOCs) in photochemical smog formation and atmospheric chemistry, it is critical to generate accurate BVOC emission inventories. Assembling such inventories requires reliable characterization of the areal coverage of plant species to quantify the leaf mass of BVOC-emitting vegetation. A recent GIS-based description of vegetation cov- erage in the natural areas of California is provided by the Gap Analysis Program (GAP) database. We conducted an assessment of this database in Central California through ground-based vegetation surveys to evaluate the use of GAP for developing a BVOC emission inventory for Central California. A modified stratified randomized sampling design was used to select and assess 18 GAP polygons. Quantitative vegetation surveys were conducted along belt transects in polygons dominated by trees and along line transects in polygons dominated by shrubs to determine percent cover of plant species for comparison to GAP data. The species listed by GAP accounted for a range of 0—88% of the relative cover in the polygons, with a mean of 43%. Of the 76 species listed by GAP for primary, secondary and tertiary assemblages for which data were collected (those species above the survey height), 33 were found to be correctly listed within their respective assemblages, 13 were found to be listed for the wrong assemblage, and 30 were below percentages of co-dominants of any assemblage. In the 18 polygons, a total of 51 additional species not listed by GAP were found to be present in amounts sufficient to consider them as potential co-dominants. Summed over all 18 polygons, BVOC emission indices based on field data were 20% less than those based on GAP, but for individual polygons differences ranged from — 100% to more than + 100%. Key Words: BVOC emission inventories require data for emission rates, areal coverage, and leaf mass of re- spective plant species. With the proposal of a tax- onomic methodology for assigning isoprene and monoterpene emission rates to unmeasured plant species (Benjamin et al. 1996), emission rates can in principle be estimated for many of the 6,000 plant species in California based on measurements within respective families and genera (Karlik and Winer 2001la; Winer and Karlik 2001). For Cali- fornia, vegetation spatial distribution and compo- sition has been described for urban and natural ar- eas within Orange County and the non-desert por- tions of Los Angeles, Riverside, and San Bernar- dino Counties (Winer et al. 1983: Miller and Winer ' jfkarlik @ucdavis.edu Vegetation survey, biogenic hydrocarbons, Gap Analysis Program, GAP, BVOC. 1984; Horie et al. 1991; Benjamin et al. 1997), and limited studies of plant composition have also been conducted for the Phoenix, AZ, urban area (Karlik and Winer 2001b) and the urban areas of Santa Bar- bara and Ventura Counties (Chinkin et al. 1996). However, a validated inventory of vegetation spe- cies composition and spatial distribution, specifi- cally to develop a BVOC emissions inventory, has not been established for the extensive areas of nat- ural vegetation in the San Joaquin Valley air basin. A potential source of information concerning vegetation in the natural areas of the Central Valley and the Sierra Nevada is the Gap Analysis Program (GAP) database, which is coordinated by the Unit- ed States Geological Service—Biological Resources Division (formerly the National Biological Service) to identify the distribution and management status of plant communities, especially to identify gaps in 100 habitats for plant or animal species needing protec- tion. GAP compiled a geographic information sys- tem (GIS) database (based primarily on remote- sensing data) describing vegetation type and dom- inance in terms of areal coverage (Davis et al. 1994, 1995). Unlike other vegetation maps which describe plant geography only in terms of plant communities, the California GAP database de- scribes vegetation in quantitative terms using dom- inant plant species and species assemblages. Because BVOC emissions inventories rely on species-specific measurements of both leaf mass and BVOC emission rates (Benjamin et al. 1997), GAP offers the advantage of providing species-spe- cific vegetation distribution data. Moreover, the GAP GIS database is recent for California (Davis et al. 1995) and provides seamless coverage of the state. Although large-area small-scale GIS databas- es based on remote-sensing data, such as GAP, offer a potentially inexpensive and relatively simple ap- proach to characterizing the distribution and species identities of natural vegetation within an airshed, use of such GIS databases for BVOC emissions in- ventory development requires evaluation of their accuracy and reliability for this purpose through ground-based observations. The present GAP validation study was modeled after a previous study conducted in San Diego County (Winer et al. 1998; Chung and Winer 1999). Quantitative vegetation surveys were con- ducted along belt transects in polygons dominated by trees, and along line transects in polygons dom- inated by shrubs, in order to determine percent cov- er of major plant species for comparison with GAP listings. We report here the results of a ground-based as- sessment of the GAP database within the Great Val- ley and Sierra Nevada ecological regions using vegetation surveys of representative GIS polygons. The surveys employed a modified stratified random sampling approach and a survey protocol based in part on the recommendations of the developers of the GAP database (Stoms et al. 1994), and refine- ments from the preceding study of GAP in San Di- ego County (Chung and Winer 1999). Data gath- ered from field surveys conducted during the 1999 and 2000 summers were used to assess the accuracy and concomitant utility of the GAP GIS database for providing quantitative information of plant spe- cies identities and coverages for BVOC emission inventories. METHODS Acquisition and Preparation of the GAP Database As noted earlier, the purpose of GAP was to identify the distribution and management status of selected components of biodiversity. The central tool of this program was an ARC/INFO GIS data- base with plant species and vegetation class attri- butes associated with polygons within a defined MADRONO [Vol. 49 geographic region. This database was generated from summer 1990 Landsat Thematic Mapper sat- ellite imagery, 1990 high altitude color infrared photography, vegetation type map surveys based on field surveys conducted between 1928 and 1940, and miscellaneous vegetation maps and ground sur- veys (Davis et al. 1995). Polygons were delimited based on climate, physiography, substrate, and dis- turbance regime. Landscape boundaries were sub- jectively determined through photointerpretation by expert personnel so that between-polygon variation was greater than within-polygon variation. The final result was a vegetation map with a 100 ha mini- mum mapping unit and a 1:100,000 mapping scale (Davis et al. 1995). The GAP database for the southern portion of the Sierra Nevada and Great Valley ecological regions was obtained at the beginning of the project. Seven hundred forty-two polygons in the Great Valley and 1420 polygons in the Sierra Nevada ecological re- gions were found within the counties of Kern, Tu- lare, Kings, Fresno, and Madera. For each polygon in the database, a primary vegetation assemblage was listed. For most polygons a secondary vege- tation assemblage was listed, and for some poly- gons a tertiary vegetation assemblage was also giv- en. The primary assemblage was defined as the as- semblage covering the majority of the polygon, and the secondary and tertiary assemblages as covering relatively smaller areas of the polygon. Each as- semblage consisted of up to three co-dominant overstory species, each with a minimum of 20% of the relative cover of that assemblage. Relative cov- er of a given plant species within an assemblage was the fraction of total assemblage vegetation cov- er occupied by the given species. Polygon Selection GAP data for each polygon were used to gener- ate an isoprene and monoterpene index for the polygon (Winer et al. 1998; Chung and Winer 1999). Although polygons with high emissions in- dices may hold more interest for emissions mod- elers, after lengthy discussions and review of com- ments on a previous study design for GAP (Winer et al. 1998; Chung and Winer 1999), we decided to use a random selection process rather than focusing on “‘high emitting”’ polygons for field validation in this study. To remain a candidate for field valida- tion, the polygon had to be below the atmospheric boundary layer, taken as 1800 m elevation, and within the San Joaquin Valley air basin. Further se- lection from the remaining polygons involved an iterative process accounting for feasibility, includ- ing physical access and permission to survey veg- etation. A road map was overlaid on the area to see if there was access by roadways, and if so a uni- versal transmercator (UTM) grid was generated. Polygons with a large public land component (e.g., within National Forest) were favored due to the rel- 2002] KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 101 TABLE 1. POLYGONS FROM THE GREAT VALLEY AND SIERRA NEVADA GAP DATABASE SELECTED FOR FIELD SURVEY. robes Approximate Area Elevation Transect Centerpt. Transect (no.) GAP No. location (ha) (m) Type (no.) (no.) 1 16541 Kern National Wildlife oper 66 Line 4 16 Refuge 2 17198 Kern River & Hwy 99 284 120 Belt 3 8 3 15485 Lemoore 256 64 Line 3 11 4 17023 Buttonwillow 254 82 Line 2 8 5 16442 Glennville 1383 1520 Belt 3 12 6 16753 Keysville 195 1570 Belt 3 12 Hy 16908 Kern River Canyon 640 760 Belt e: 12, 8 14572 Sequoia National Park 4431 1280 Belt 3 12 9 16791 Bodfish 309 1200 Belt 3 12 10 16273 California Hot Springs 2688 1570 Belt 4 14 11 13790 Sequoia National Park 635 1270 Belt 2 8 12 13795 Kings Canyon 2473 1340 Belt 1 2 13 16269 California Hot Springs 341 930 Belt 2 6 14 16687 Lake Isabella 1019 845 Belt 3 f2 15 16783 Alta Sierra 476 1060 Belt 2, 8 16 16776 Alta Sierra 642 1340 Belt 2 4 17 16756 Alta Sierra 1969 1080 Belt 2 4 18 16735 Alta Sierra 812 1030 Belt 2 4 ative ease of gaining permission to conduct surveys on such properties compared to privately owned properties. Based on these criteria and the time and resourc- es available for this research, 18 polygons in south central California were selected and surveyed for the present study, as seen in Table 1. Four polygons (nos. 1—4), located on the Central Valley floor, con- sisted primarily of shrubs and herbaceous vegeta- tion, and fourteen polygons (nos. 5—18), located in the Sierra Nevada mountains and foothills, consist- ed primarily of woodland and forest vegetation thie. ft): A subsample of polygons was selected as a test for correctness of the geographic location of a spe- cific GAP polygon; in other words, a test of the registration of the GAP database. Three surround- ing polygons adjacent to polygon 15 were selected for survey during the summer, 2000, sampling sea- son. The plants found in these surrounding poly- gons could then be compared with those listed for the center polygon to see if plant communities list- ed for the center polygon were found instead in a surrounding polygon. The three surrounding poly- gons were no. 16 located to the southeast, no. 17 located on the western end, and no. 18 located to the northeast of polygon 15. Selection of Sample Elements If permission was obtained to access most of a polygon, sample elements of 500 * 500 m squares were selected by overlaying a 500 X 500 m UTM grid on the polygon, assigning sequential numbers to every grid square within | km of a road, and randomly selecting the centerpoint locations for the needed number of elements. The number of center- points and corresponding elements varied with polygon size. For polygon areas of <1000, 1000— 10,000, and >10,000 ha, two, three, and four cen- terpoints were chosen, respectively, although ter- rain or accessibility sometimes limited the number of centerpoints. This method was similar to the one employed in the Utah GAP validation project (Ed- wards et al. 1995) and that of Chung and Winer in San Diego County (1999). In several cases, suitable survey sites were not available within the vicinity of a road, so hikes of up to two hours along a trail were needed to reach the desired area within the polygon. Vegetation Survey Protocol The specific survey protocol chosen depended on the type of vegetation being assessed. Within the polygons dominated by trees, surveys were per- formed by a team of two along 6 m wide, 500 m long belt transects orthogonal at the centerpoint in most elements. Six meter wide belt transects make the mechanics of sampling easier while not signif- icantly compromising accuracy (Lindsey 1955). For these belt transects, the surveyors walked 250 m north, south, east, and west away from the center- point, using a magnetic compass to maintain course. Within polygons dominated by shrubs, the sur- vey for each element consisted of two 300 m line transects orthogonal at the centerpoint. Line tran- sects have been used to estimate relative cover for chaparral (Bauer 1943) and for sage scrub (Kent and Coker 1992; Zippin and Vanderwier 1994). The minimum square-shaped area needed to encompass a sample element within a polygon was therefore 25 ha for forests and woodlands, and nine ha for scrub and chaparral. The survey team located the centerpoint of a par- 102 MADRONO Location of Great Valley polygons Nos. 1-4 Fic. 1. ticular sample element using a global positioning receiver (GPS) locked onto UTM coordinates gath- ered from the GAP database. A handheld GPS unit (Garmin 12XL), with an accuracy of approximately 22115 im Gr 22D mn, wa ISLS) ancl AVOO, mesoecinvelby, was employed. Plant community and site descrip- tions were recorded and elevation at the centerpoint was determined using a hand-held altimeter (Pretel Instruments). Data Collection For belt transects, data collected included the crown radii, crown height, and diameter at breast height of trees and the crown dimensions of shrubs. For line transects, plant species identity, crown height and number of 0.1 m segments along a meter tape occupied by that plant species were noted. Plants such as grasses and forbs below a height of about 0.6 m were not recorded. Additional details of plant measurement methods have been reported previously (Winer et al. 1998; Chung and Winer 1999). Plant nomenclature follows Mabberley Geom: [Vol. 49 Location of Great Valley polygons Nos. 5-18 Locations of the polygons selected for field survey from the GAP database. Data Analysis Data analysis followed the example of Chung and Winer (1999). For each polygon, the GAP da- tabase listed primary, secondary, and sometimes tertiary species assemblages and the estimated areal proportion (p) of each assemblage within a poly- gon. Each species in a listed assemblage was a co- dominant, providing =20% relative cover within the assemblage. Therefore, the expected coverage of any species listed in the GAP database for a given polygon was =0.2p. For example, in polygon 5 Quercus kelloggi was listed as a co-dominant in a primary assemblage that occupied 60—70% of the polygon. Using a mean value of 65% for p, GAP predicted Quercus kelloggi would cover 20.2 X 65%, or 213% of the polygon. The polygon coverage of plant species inferred from the GAP database by this procedure was com- pared with the cover data gathered from the field surveys in the 18 selected polygons. First, the cov- erage of each species within each sample element of a polygon was calculated. Then from the species coverage for each sample element, the mean cov- 2002] erage and upper limit of the two standard error (SE) confidence intervals for the polygon were calculat- ed, corresponding to an 85% confidence interval (McClave and Dietrich 1985). RESULTS Species Composition and Abundance within GAP Polygons Table 2 summarizes results for the 18 polygons surveyed, listing the most abundant species ob- served for each polygon, the percent abundance predicted from the GAP database, the percent abun- dance determined by the field surveys, and the up- per limits of a two SE interval of the percent com- position. Plant species not listed by GAP or not found in surveys at =1% are omitted from this ta- ble, but may be found in the report of Winer and Karlik (2001). Total plant cover within the polygon ranged from as little as 7%, as found in polygons 2 and 9, up to 82% as found in polygon 12. In general, most of the sample cover in a polygon was attributable to a few species and many of the most abundant species found within the polygon were listed as co-dominants by the GAP database. How- ever, the percentages of these GAP co-dominant species varied greatly. Total cover of GAP co-dom- inant species cover ranged from as little as 0% as found in polygon 18 up to 66% as found in polygon 2: Relative cover of GAP-listed species compared to sampled species can be derived from the per- centages listed in Table 2. For the Great Valley polygons (nos. 1—4), GAP species listings ranged from 29-72% of the plant species found. The sum of GAP species percentages for these polygons was 46, and for all species sampled, 100. Thus, GAP- listed plants accounted for 46% of the relative cov- er for polygons nos. 1—4 considered together. For the Sierra Nevada polygons (nos. 5—18), the rela- tive cover of GAP-listed species for each polygon ranged from O—88%, and the relative cover was 43% for polygons nos. 5—18 considered together. If all 18 polygons were considered together, GAP-list- ed plants accounted for 43% of the relative cover overall. To investigate whether GAP listings were more accurate for large vs smaller polygons, poly- gon size was multiplied by the corresponding per- centages found for GAP-listed species vs for all species. The results suggested polygon size did not influence accuracy of GAP listings, since relative cover of GAP species was then found also to be 43%. The observed sample cover of some co-domi- nants in GAP polygons often substantially exceed- ed the minimum predicted values. For example, in polygon 12 Quercus chrysolepis provided 51% of the polygon sample cover though =7% and =3% were predicted from the GAP listings for the sec- ondary and tertiary assemblages, respectively. In polygon 5 Quercus kelloggii provided 31% of the KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 103 polygon sample cover when =13% was predicted by GAP. In contrast, several polygons possessed co-dom- inant species that were found to be well under the predicted GAP percentages. For example, polygon 9 was found to have 1% mean sample cover of Yucca whipplei when GAP predicted the species to have 211% and =7% sample cover for the primary and secondary assemblages, respectively. In poly- gon 12, Pinus ponderosa was predicted by GAP to be found with sample cover =11% when the field study found it to be only 0.3% sample cover. Because Quercus may be the most important ge- nus of native woody plants with regard to BVOC emissions in California’s airsheds, we compared the percentage of all oak species sampled to that in- ferred from GAP listings. For the 14 Sierra Nevada polygons (nos. 5—18), the mean coverage per poly- gon of oaks calculated from GAP listings was 17%, while from our field surveys oaks averaged 29% of the total cover. For all 18 polygons, the mean cov- erage of oaks calculated from GAP per polygon was 13% and from field surveys 22%. Since GAP listings give a lower limit for species abundances, one might expect to find oaks at the same or higher percentages than GAP listings. Considering poly- gons individually, for six of the 18 polygons we found oak species percentages within a factor of two of the respective GAP-predicted percentages. For an additional five polygons, oaks were listed as comprising zero percent cover, and were not found in the field surveys, in agreement with GAP list- ings. Thus, for 11 of 18 polygons, GAP listings for oaks were in reasonably good agreement with field data. For three polygons, nos. 8, 14, and 18, oaks were found at 20, 4, and 43% although no oak spe- cies were listed by GAP. Thus, overall GAP data were in reasonable agreement with field data, and the lower limit of oak coverages as given by GAP may underestimate oak abundance. Since Quercus species vary over almost two or- ders of magnitude in isoprene emission rate (Csiky and Seufert 1999), the accuracy of oak species list- ings for any vegetation database is important for BVOC emission inventories. Using the data in Ta- ble 2, we considered oak species accuracy for each polygon by calculating the sum of absolute values of field percentage found for each oak species mi- nus the respective percentage inferred from its GAP listing. These values for polygons ranged from 0% for the polygons where no oaks were listed or found to more than 40% for polygons 12, 15, 17, and 18 with a mean for all polygons of 19%. The accuracy of oak species listings did not appear to be related to polygon size. We also evaluated the data in Table 2 with regard to non-oak genera and species considered to be moderate or high BVOC emitters, specifically those thought to have emission rates greater or equal to 10 or 2 pg g-' h'' for isoprene and monoterpenes, respectively, as based on measured values or tax- 104 MADRONO [Vol. 49 TABLE 2. MEASURED SPECIES COVER COMPOSITION OBSERVED IN SAMPLED GAP POLYGONS LISTED IN ORDER OF OBSERVED SAMPLED COVER. Species with mean sampled cover of <1% or not listed in GAP are omitted. N.D. = no data. For example, for Avena spp. and Bromus spp. species were observed but below survey height (about 0.6 m), and data were not recorded. Sampled cover GAP predicted Mean sampled (s + 2SE) Polygon Species cover (%) cover (%) (%) 1 Cyperus difformis — L/ 30 Typha spp. =11 & =3 15 38 Scirpus acutus —- 6 17 Xanthium strumarium — 4 8 Brassica nigra — 3 7 Distichlis spicata — 3 6 Scirpus californicus — 1 3 Baccharis salicifolia _ 1 2 Suaeda ramossissima —_ 1 2 Atriplex coronata — 1 2 Allenrolfea occidentalis 27] 1 22 Salix spp. =] 0.2 0.6 Carex spp. 22\lil 6 228 0.0 — Juncus spp. == 0.0 — Tamarix spp. == WSO =3 0.0 — Total of sample cover 3S) GAP co-dominants 16 2 Unknown #1 — 3 7 Populus fremontii = D 3 Platanus racemosa -— i D; Salix spp. — 1 2 Distichlis spicata 21/3) eG, ei) N.D. — Total of sample cover 7 GAP co-dominants 2 3 Allenrolfea occidentalis 22117 Go 223 21 43 Salix sp. — 9 Da Populus fremontii = D, 5 Eucalyptus spp. =3 0.0 = Total of sample cover 32 GAP co-dominants 3 4 Atriplex polycarpa 22 (9) 5) 9 Adenostoma fasciculatum — 2 6 Avena spp. and Bromus spp. 220) N.D. — Total of sample cover 8 GAP co-dominants 5 5 Quercus kelloggii =13 Syl 4] Quercus wislizenii 2213) 6's 227/ 14 19 Quercus garryana =] 6 sI97/ Calocedrus decurrens _- =) 9 Cercocarpus betuloides =|) 5 8 Abies concolor 2 5) Pinus ponderosa =13 1 2, Quercus berberidifolia ] 1 Total of sample cover 65 GAP co-dominants Si 6 Quercus kelloggii NB ce 227) 14 ES) Pinus ponderosa 22/3) (6 227) 6 13 Arctostaphylos spp. — =) 11 Quercus chrysolepis — 4 10 Abies magnifica —- 3 6 Calocedrus decurrens — 2 7 Abies concolor =13 1 3 Cercocarpus betuloides — ] 2 Quercus wislizenii =7 ] 1 Total of sample cover 38 GAP co-dominants DD, 2002] Polygon I 10 11 KARLIK ET AL Species Quercus wislizenii Quercus douglasii Rhamus crocea Ceanothus cuneatus Quercus garryana Platanus racemosa Pinus sabiniana Quercus dumosa Aesculus californica Adenostoma fasciculatum .. GAP ASSESSMENT IN CENTRAL CALIFORNIA TABLE 2. CONTINUED. GAP predicted cover (%) = 7/ Avena spp. and Bromus spp. =11 Total of sample cover GAP co-dominants Abies concolor Quercus douglasii Quercus kelloggii Calocedrus decurrens Ceanothus integerrimus Aesculus californica Quercus chrysolepis Pinus ponderosa Pinus lambertiana Cornus nuttallii Sequoiadendron gigantea Umbellularia californica Adenostoma fasciculatum Total of sample cover GAP co-dominants Chrysothamnus nauseosus Juniperus californica Yucca whipplei Eriogonum fasciculatum Artemisia tridentata Total of sample cover GAP co-dominants Quercus douglasii Aesculus californica Quercus wislizenii Quercus kelloggii Quercus garryana Quercus chrysolepis Cercocarpus betuloides Pinus sabiniana Rhus diversiloba Ceanothus cuneatus Ribes sp. Total of sample cover GAP co-dominants Cercocarpus betuloides Ceanothus integerrimus Ceanothus cuneatus Quercus dumosa Aesculus californica Quercus chrysolepis Arctostaphylos sp. Umbellularia californica Arctostaphylos mewukka Total of sample cover GAP co-dominants Mean sampled cover (%) io) ONFRRFN WW ORR KYPNWWHW HK OW WO oo ~— 105 Sampled cover (S—Z2SB) (%) Wore BWW BND WW © 106 Polygon 12 13 IS) 16 7 Species Quercus chrysolepis Calocedrus decurrens Umbellularia californica Cercocarpus betuloides Ceanothus cuneatus Quercus dumosa Arctostaphylos sp. Salix spp. Pinus ponderosa Abies concolor Aesculus californica Pinus contorta Total of sample cover GAP co-dominants Quercus wislizenii Aesculus californica Quercus douglasii Ceanothus cuneatus Quercus dumosa Umbellularia californica Fremontodendron californicum Cercocarpus betuloides Pinus sabiniana Avena spp. and Bromus spp. Total of sample cover GAP co-dominants Ceanothus cuneatus Pinus sabiniana Quercus douglasii Quercus wislizenii Quercus dumosa Ephedra california Mimulus aurantiacus Yucca whipplei Adenostoma fasciculatum Juniperus californica Artemisia tridentata Avena spp. and Bromus spp. Total of sample cover GAP co-dominants Quercus kelloggi Quercus douglasii Quercus garryana Quercus dumosa Aesculus californica Quercus wislizenii Ceanothus cuneatus Pinus sabiniana Cercocarpus betuloides Pinus ponderosa Avena spp. and Bromus spp. Total of sample cover GAP co-dominants Ceanothus cuneatus Quercus lobata Quercus douglasii Aesculus californica Quercus wislizenii Quercus kelloggii Quercus garryana MADRONO TABLE 2. CONTINUED. GAP predicted cover (%) =7 & =3 =11 Mean sampled cover (%) [Vol. 49 Sampled cover (s + 2SE) (%) 2002] KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 107 TABLE 2. CONTINUED. Polygon Species Quercus dumosa Ribes sp. Pinus sabiniana Cercocarpus betuloides Avena spp. and Bromus spp. Total of sample cover GAP co-dominants 18 Quercus douglasii Quercus wislizenii Aesculus californica Ceanothus cuneatus Quercus dumosa Pinus sabiniana Cercocarpus betuloides Adenostoma fasciculatum Unidentified chaparral shrubs Bare exposed rocks Total of sample cover GAP co-dominants onomy (Benjamin et al. 1996; Karlik and Winer 2001a). For the 11 polygons where such genera or species were listed, the field surveys gave percent- ages lower than those inferred from GAP for seven polygons, and higher than GAP for four polygons. For the seven polygons where no genera or species with moderate or high emission rates were listed, the field surveys found an absence of such genera or species in two polygons, and for the other five polygons percentages of emitting species ranged from 1—11% of total area. In considering the mean of GAP listings for BVOC-emitting species vs the mean of the field survey data for all 18 polygons, GAP predicted a mean coverage per polygon of 9% for these species, and field data were in good agree- ment with a mean of 8% of such species found. Thus, on average GAP data were harmonious over- all with field observations for non-oak genera and species considered to be important BVOC emitters. Correctness of GAP Listed Species within Species Assemblages Species found within the polygons in the field were compared to their GAP listings and assessed for correct placement based on assemblage data, and the results are given in Table 3. A species was considered to be correctly listed when the percent- age found in the field within two standard errors (s + 2SE, Table 2) exceeded the GAP-predicted per- centage. A species was considered listed incorrectly when listed by GAP as a co-dominant in a partic- ular assemblage (primary, secondary, and tertiary) but found at a lower percentage cover so as to place it within a different assemblage. Potential co-dom- inant species were defined as species not listed by Sampled cover GAP predicted Mean sampled (S)-- 2SB) cover (%) cover (%) (%) — 4 6 — 2 6 — p? 6 — 1 1 =>3 N.D. — 62 8 = 25 40 — 15 40 — 9 Dal — 9 9 — 3 9 — 1 BY — 1 2 =13 0.0 — =5 N.D — >3 N.D — 63 0) GAP as present in the polygon but found in field surveys to have cover percentage large enough to at least fall within the tertiary assemblage of a par- ticular polygon. When GAP listed no species for the secondary or tertiary assemblage, an arbitrary value of =7% and =3% up to the next greater listed assemblage percentage were assigned, respectively, to identify potential species belonging to a partic- ular assemblage. We note that GAP assigns mini- mum percentages to plant species coverages, but does not assign maxima; therefore, species listed in two or more coverage classes were considered to have a correct listing in these classes if present in sufficient quantity for the greater percentage re- quirement. For example, in polygon 6 the species Quercus kelloggii was listed in both primary and secondary assemblages, found to be present at 14%, and considered to be correctly listed for both assemblages. The agreement of field results with GAP data varied among polygons, as seen in Table 3. There were several polygons (nos. 3, 5, 6, 10, 13) in which the field results agreed with all or the ma- jority of GAP listings. In contrast, there were sev- eral instances where species listed by GAP in either the primary, secondary, or tertiary assemblage were not observed in the polygon in sufficient abundance for for their respective assemblage. For example, in polygon 6 Abies concolor in the primary assem- blage and in polygon 7 Quercus douglasii in the secondary assemblage were found with coverage percentages below those for their respective assem- blages. Potential co-dominant species were noted in several polygons, such as in polygons 8 and 17, where seven species were found in percentages that [Vol. 49 MADRONO 108 13830]]2Y SNIAINOE psosapuod snuig 118.30]]99 SNILINO psosapuod snuig QUON AOJOIUOD Sa1gqy AOJOIUOD Saiqy a 9 (CG) SualINIap SNAPAIOIDI oUON Tt, NUAZ1I]SIM SNILINO UAZISIM SNILINO DUDKAIDS. 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SN]NISAV 50 cm diameter breast height [dbh]) California black oak and ponderosa pine (Pi- nus ponderosa). Other less dominant tree species included interior live oak (Q. wislizeni), Douglas- fir (Pseudotsuga menziesii), white fir (Abies con- color), sugar pine (P. lambertiana), and incense ce- dar (Calocedrus decurrens). Seedling and sapling California black oak and ponderosa pine dominated the subcanopy at one stand, while the other stands had little subcanopy. The shrub layer was generally sparse, and deerbrush (Ceanothus integerrimus) and manzanita (Arctostaphylos spp.) were the most common shrubs. The herbaceous layer was domi- nated by a sparse to dense cover of mountain mis- ery (Chamaebatia foliolosa) and seedlings of Cal- ifornia black oak. California black oak was rela- tively widespread throughout the study area and oc- curred in stands with trees of all ages with varying amounts of conifers. The study stands were repre- sentative of those with mature, large diameter Cal- ifornia black oak in the central Sierra Nevada, al- though these stands are rare in this area (Garrison et al. 1998). METHODS Seedling densities and associated vegetative characteristics were sampled from 30 0.04-ha plots in each stand (known as Stands 1—4). Each stand had two adjacent 10.6-ha subplots, and 15 0.04-ha circular plots were randomly selected and measured in each subplot at intersection points of a 25-m by 25-m grid. To ensure that the entire subplot was sampled, one circular plot was randomly located on each of 11 transects in the grid and sampled in July—August 1994. Four additional circular plots were selected from the same transects and sampled in July-August 1995 for a total of 15 circular plots per subplot and 30 circular plots per stand. Seedling and understory vegetation data were measured at all circular plots in 1995. Data were taken on di- MADRONO [Vol. 49 ameter (cm) and heights (m) of trees, cover (%) of the overstory canopy and understory, and stem den- sities (number/ha) and basal areas (m7/ha) of all live stems 212.7 cm dbh and =2 m tall. Seedlings (oaks <2.5 cm basal diameter) were counted from five 1- m? circular plots located at the 0.04-ha circular plot center and 5 m away from the plot center along axes pointing North (0°), East (90°), South (180°), and West (270°). Data were also collected on sap- lings (oaks 2.5—15 cm basal diameter and <2 m tall), but saplings occurred on 8% (10 of 120) of the vegetation plots so they were too rare to be analyzed. Because most data did not meet assumptions and distribution requirements for parametric statistics, the data were transformed (Zar 1996). Log trans- formations (log,) + 1.0) (Zar 1996) were used for averages for seedling counts, basal area, stem den- sity, heights and stem diameters for all trees, Cali- fornia black oak, and conifers. Cover for understo- ry and overstory canopy were given arcsine trans- formations [arcsine radian degrees (square root (proportion canopy cover + 1))] (Zar 1996). Analysis of variance with Bonferroni pairwise comparisons were used to determine differences among stands for the vegetation attributes. Pearson correlation coefficients for the transformed data was used to determine preliminary relationships be- tween vegetation attributes and seedling densities with data from all stands combined and separately for Stand 1. While combining data from all stands for the correlations is considered pseudoreplication (Hurlbert 1984), the correlations were for explor- atory purposes to select a smaller subset of habitat variables for more detailed analyses and no statis- tical inferences were made using these results. Vegetation attributes with statistically significant (P < 0.05) correlations were used in a backward stepwise general linear model to determine stand differences in seedling densities and which vege- tation attributes had the greatest effect on seedling density. A multiple linear regression analysis was conducted for Stand 1 because it had the greatest seedling densities and frequency of plots with seed- lings so it provided an opportunity to assess how habitat attributes influence seedling densities at the stand level. Understory cover measurements were not included in the general linear model or regres- sion because two of the five understory attributes were highly correlated (P < 0.02) with seedling density; this correlation indicated data interdepen- dence and redundancy that biased relationships be- tween the tree layer and seedlings. Statistical anal- yses were conducted using SYSTAT (SPSS Incor- porated 1999). Summary statistics and scatterplots for each stand were used to assist interpretation of the general linear model results. Throughout this paper, the term “‘oak”’ is synonymous with Califor- nia black oak as almost 100% of the tree oaks and 100% of the oak seedlings and saplings measured were California black oak. 2002] RESULTS Stand Attributes The four stands consisted of relatively large di- ameter California black oaks and smaller diameter conifers with closed canopies, large amounts of basal area and moderate stem densities (Table 1). Sixteen of the 21 (76%) vegetation attributes mea- sured differed (P < 0.024) among the four stands. Stand 1 had greater oak seedling densities (#/ha), greater conifer tree densities (#/ha), greater oak un- derstory cover (%), lesser oak tree densities (#/ha), smaller diameters (cm) of conifer trees, and shorter conifer trees (m) than the other stands (P < 0.001). Stem densities, understory and overstory cover, tree diameters, and tree heights were the attributes that mostly differed among the stands (P < 0.024) (Ta- ble 1). Basal area (m7/ha) of all trees and oak were similar (P > 0.197) among stands. At all stands, 63-90% of the sample plots had at least one oak seedling indicating that some seedlings occurred over much of the area within and among stands (Fig. 1). Stands were at similar elevations (1220— 1320 m); slopes varied from 5—30%, and aspects included E, SE, S, and W (Table 1). Habitat Attributes Affecting Seedling Densities Of the 20 vegetation attributes analyzed for their possible effects on seedling densities at the four stands, seven (35%) attributes were correlated (n = 95-120; P < 0.038) with seedling density. Five of these seven attributes were from the tree layer (co- nifer tree density [#/ha], oak basal area [m?/ha], av- erage diameter [cm] of oaks and conifers, and av- erage height [m] of oaks) and were used in the gen- eral linear model. The other two attributes (% total understory cover and % oak understory cover) ere not included in the general linear model because they are highly correlated (n = 120, P < 0.02) and redundant variables for seedling density. At Stand 1, four (20%) of the 20 vegetation attributes (% oak and conifer overstory cover, oak basal area [m2?/ha], and % oak understory cover) were correlated (n = 30, P < 0.049) with seedling density. Average diameter (cm) of California black oak and stand were only attributes retained in the gen- eral linear model (Table 2). Seedling densities dif- fered (P < 0.008) among stands and increased (P < 0.001) as average oak tree diameter increased (Table 2, Fig. 2). This relationship was due some- what to the greater number of seedlings and larger diameter trees at Stand | (Table 1, Figs. 1 and 2). With Stand 1, seedling density increased with in- creasing oak basal area (P < 0.028). DISCUSSION California black oak seedlings were prevalent at one of four stands in the central Sierra Nevada, and the stands differed statistically in many vegetation attributes including densities, diameters, and GARRISON ET AL.: CALIFORNIA BLACK OAK SEEDLINGS 117 heights of trees. Oak seedling densities differed among stands and increased with increasing aver- age diameters of the surrounding oak trees. More seedlings under larger diameter trees are expected because larger diameter California black oaks are older (Garrison et al. 2002) and acorn production increases with increasing age and diameter of Cal- ifornia black oak (McDonald 1969; Garrison et al. in press). Older trees also have longer periods of time to deposit acorns on the ground that germinate into seedlings so greater numbers of seedlings ac- cumulate under larger trees. All stands were dominated by California black oak and had equivalent total overstory canopies, but slopes, aspects, and oak overstory canopy cover varied somewhat among stands. Presence of Cali- fornia black oak seedlings is negatively associated with solar radiation and positively associated with canopy cover of California black oak (Standiford et al. 1997), and probabilities of seedling occur- rence at the four study stands were between 80— 90% using the graphs from Standiford et al. (1997). Stand 1 had the greatest number of seedlings and the least amount of oak overstory cover (36%) as well as the least amount of solar radiation due to a relatively flat slope. Furthermore, seedling densities increased with increasing oak basal area in Stand 1. Although seedling densities increased with in- creasing oak diameter in plots from all four stands, oak basal area was also correlated (r = 0.194, n = 120, P < 0.034) indicating that both vegetation at- tributes have affect seedlings. Seedling presence and abundance varies across small and large geographic areas as seedlings oc- curred on 63—90% of our survey plots in the central Sierra Nevada yet densities varied four to fivefold among stands. Other studies reported California black oak seedlings present on 83% of survey plots in the southern Sierra Nevada (Standiford et al. 1997) and 62% of sample plots in the range of the California black oak habitat in California (Bolsin- ger 1988). Seedlings are germinated acorns, so acorn pro- duction by parent trees ultimately affects seedling densities. California black oak acorn production has considerable spatial and temporal variation (Koenig et al. 1994) so acorns are limited at certain sites and particular times. In our stands, plots with larger oaks had lower stem densities with varying amounts of canopy cover so germinating acorns and seedlings received varying amounts of sunlight. Acorns deposited in areas with lesser canopy cover germinate and grow at greater levels that acorns in areas with greater canopy cover (Savage 1994; Standiford et al. 1997), and this pattern occurred at Stand 1 where seedling densities were greatest at moderate (15-55%) canopy coverages. Seedling re- generation of California black oak is clumpy as seedlings concentrate around drip lines of parent trees (Savage 1994; McDonald and Tappeiner 1996). Furthermore, acorns are distributed and [Vol. 49 700'0 ely 9100 09¢ 5000 erp 100°0 OL 8000 Il'y €00°0 6L' L950 g9°0 100°0 IL'€l 100°0 C8'se 610°0 Lye rL0'0 LES © Z 100'0 fia go v70'0 Woe al gor'0 L6°0 < 2 100°0 Wate 100°0 99'L 100'0 LS'8 L100 Ese r8L0 8€0 L610 gol 100'0 E091 onjea-g A \VAONV 118 Of Oc al S) OcEl OCCI alc + Sc de + 897 del + srl VVI+Cc0C LI + Vol all + Le q9r + Ler a€9¢o¢+91¢ sl GG = ih iis ec + C6PV LOE+9LE ace + S 8p 00 + 00 c0 + €0 Gd80+ C7¢ a8c + 8S aoo0+ el €@ZL0+ LI €ve+ cle DE +VLI i) @ ae (I tete Le+vle qa9e + SL? 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GARRISON ET AL.: CALIFORNIA BLACK OAK SEEDLINGS 119 Stand 2 29 20 19 es 10 Number of plots i 12345 6 7 8 9 1011 12 131415 16 1718 Seedlings per sq. m Stand 4 Number of plots Oo 1 2 3 4567 8 9 101112131415 1617 18 Seedlings per sq.m Frequencies of plots with varying densities of California black oak (Quercus kelloggii) seedlings (number/m7?) measured in 1995 at four 21.1-ha study stands in Placer County, California. The number of plots without seedlings in the 0—1 seedlings/m? density class were Stand 1 = 3, Stand 2 = 8, Stand 3 = 4, and Stand 4 = 11. cached by western gray squirrels (Sciurus griseus) (McDonald 1969) and Steller’s jay (Cyanocitta stel- leri) (personal observation, Fuchs et al. 1997). Many cached acorns are not consumed by wildlife and then germinate into seedlings. In undisturbed settings, regeneration of Califor- nia black oak from germinated acorns occurs as a steady accumulation of seedlings rather than large pulses (McDonald and Tappeiner 1996), although seedlings must be aged to confirm this. Because they grow slowly under the canopy of overstory trees, oak seedlings generally have similar heights although ages can be different (McDonald and Tap- peiner 1996). Seedling accumulations were obvious at Stand 1, but seedling ages were not determined to confirm whether they had accumulated over many years or resulted from a pulse of seedlings coincident with a bumper acorn crop. One good acorn crop, however, coincident with proper site conditions may result in a single event of great 120 MADRONO [Vol. 49 TABLE 2. RESULTS OF BACKWARD STEPWISE GENERAL LINEAR MODEL ANALYSIS ON THE DENSITY OF CALIFORNIA BLACK OAK SEEDLINGS (NUMBER/m?2) AND VEGETATION ATTRIBUTES AT FOUR 21.1-HA STANDS IN PLACER COUNTY, CALIFORNIA. 2 Attributes log, transformed. © Attributes dropped from model if P-value > 0.05. Oak diameter was the covariate. ance df F Std. coeffi- Attributes Coefficient SE cient Retained in model? Stand = — — Oak dbh> 0.46 0.17 O25 Partial Correla- Dropped from model? tions Oak basal area? 0.09 = — Conifer dbh> —0.07 — — Conifer trees> —0.02 — — Conifer height’ —0.01 —- — Analysis of Covariance SS Stand 3.37 Oak diameter® 0.70 Error 7.50 acorn germination and increase seedling densities. Acorn germination, however, is considered less sig- nificant as a recruitment source than sprouting by McDonald (1969, 1990) and McDonald and Tap- peiner (1996), yet Savage (1994) expected seed- lings to be a major source of recruitment of Cali- fornia black oak in the San Jacinto Mountains. Seedlings, rather than sprouts, appear to be the ma- jor recruitment source in the four study stands be- cause approximately 70% of the mature oak trees sampled were single stems (Garrison et al. 2002). Seedling densities increased with increasing oak ES 1.0 0.5 CA black oak seedlings (#/sq. m) (log10) 0.0 Ve ere odd ee Ake) U8) 20 21 CA black oak diameter (cm) (log10) Fic. 2. Scatterplot of California black oak seedlings/m? and California black oak diameter (cm) from four 21.1-ha study stands measured in 1994 and 1995 in Placer County, California. Toler- P-value R2 0.59 3 11.48 0.000 0.40 0.89 1 Tee) 0.008 0.63 1 0.67 0.415 0.74 1 0.36 0.549 0.78 1 0.04 0.848 0.85 1 0.00 0.960 MS df F P-value R?2 1.13 3 15.46 0.000 0.41 0.70 1 9.55 0.003 0.07 103 diameter (P < 0.001) at all stands and increasing oak basal area at Stand | so diverse stand condi- tions with lesser and greater tree sizes and ages and basal areas should result in variable seedling den- sities. Larger diameter oaks have more wildlife habitat attributes such as dead branches, mistletoe (Phoradendron villosum), and acorns (Garrison et al. in press) so retaining large diameter trees is rec- ommended where land management activities might affect these trees. Acorn production varies across California black oak diameters from 35—115 cm (Garrison et al. in press) but larger trees pro- duce more acorns so retaining larger trees will maintain acorn crops and provide additional wild- life habitat benefits. Stand | received a prescribed fire on 21 October 1981, while the other stands had not burned within the last two decades. This prescribed fire and the larger diameter oaks, flater slope, and moderate oak overstory canopy cover were the most significant factors distinguishing Stand 1 from the other three stands and remains the most plausible reason why Stand | had more seedlings than the other stands. Prescribed fire appears to be a viable management tool in stands dominated by California black oak to promote seedlings and eliminate leaf litter (Ander- son 1993) as well as to reduce threats from more severe, stand-replacing fires. Prescribed fires con- ducted in late fall or early spring with low to mod- erate fuel loads caused relatively low levels of mor- tality to California black oak seedlings and sprouts (Kauffman and Martin 1990), so prescribed fires can be timed to promote seedling establishment from acorns, maintain existing seedling and sapling numbers, and damage or kill competing vegetation. Fall burns conducted prior to the drop of acorns and deciduous leaves might be the best time to burn based on our limited data and lack of assessment 2002] of the effects of prescribed fire on seedlings in our study area. California black oak grows throughout its range in many different environmental conditions, and stands occur in even and uneven-aged conditions with varying amounts of conifers and other hard- woods (McDonald 1969, 1990; Garrison et al. 2002) so management objectives and actions will differ accordingly. Retaining acorn-producing trees while opening the canopy to moderate levels through thinning could produce conditions suitable for acorn germination and seedling growth. Thin- ning oak stands also improves acorn production (Healy 1997; Standiford et al. 2000). Finally, this study was done on a limited number of sites (four) in the central Sierra Nevada with mature California black oaks so these results may have limited appli- cation to stands with different vegetative attributes. ACKNOWLEDGMENTS Ray Brumitt, Karen Durand, Catherine Fowler, Linda Louie, Jeanette Mar, Christopher Otahal, David Pratt, and Patty Sterling deserve special appreciation for their hard work sampling vegetation. Bob Heald, Doug McCreary, Phil McDonald, and Richard Standiford provided advice, information, and insights into the ecology and manage- ment of California black oak. Doug McCreary, Phil McDonald, Kristina Schierenbeck, and an anonymous re- viewer commented on earlier versions of this paper and gave helpful suggestions. This study was funded by the Deer Herd Plan Implementation Program of the California Department of Fish and Game and U.S. Forest Service. We thank Ken Mayer, Terry Mansfield, Eric Loft, Sonke Mastrup, Russ Mohr, Joelle Buffa, Susan Sharpley-Evans, Linda Tatum, Sheryl] Ducummon, Richard Johnson, Ray- mond LaBoa, Lisa Kreuger, Jeff Finn, and Ron Bertram for their interest, support, and assistance with this study. LITERATURE CITED ANDERSON, M. K. 1993. The mountains smell like fire. Fremontia 21:15—20. BOLSINGER, C. L. 1988. The hardwoods of California’s timberlands, woodlands, and savannas. USDA Forest Service Resource Bulletin PNW-RB-148. Fucus, M. A., P. G. KRANNITZ, A. S. HARESTAD, AND FE L. BUNNELL. 1997. Seeds that fly on feathered wings: acorn dispersal by Steller’s Jays. Pp. 648—650 in Pro- ceedings of a Symposium on Oak Woodlands: Ecol- ogy, Management, and Urban Interface Issues. USDA Forest Service General Technical Report PSW-GTR- 160. GARRISON, B. A., R. L. WacuHs, T. A. GILES, AND M. L. Triccs. 1998. Progress report: wildlife populations and habitat attributes of montane hardwood-conifer habitat in the central Sierra Nevada. State of Califor- nia, Department of Fish and Game, Wildlife and In- land Fisheries Division, Administrative Report 1998-1. , C. D. OTAHAL, AND M. L. TricGs. 2002. Age structure and growth of California black oak (Quer- GARRISON ET AL.: CALIFORNIA BLACK OAK SEEDLINGS 2 cus kelloggii) in the central Sierra Nevada. Pp. 665— 679 in Proceedings of the Fifth Symposium on Oak Woodlands: Oaks in California’s Changing Land- scape. USDA Forest Service General Technical Re- port PSW-GTR-184. , R. L. WaAcus, T. A. GILES, AND M. L. Triaccs. In press. Dead branches and other wildlife resources on California black oak (Quercus kelloggii). Proceedings of a Symposium on Dead Wood in Western Forests. USDA Forest Service General Technical Report. GRIFFIN, J. R. AND W. B. CRITCHFIELD. 1972. The distri- bution of forest trees in California. USDA Forest Ser- vice Research Paper PSW-82/1972. HEALY, W. M. 1997. Thinning New England oak stands to enhance acorn production. Northern Journal of Ap- plied Forestry 14:152—156. HURLBERT, S. H. 1984. Pseudoreplication and the design of ecological field experiments. Ecological Mono- graphs 54:187—211. KAUFFMAN, J. B. AND R. E. MARTIN. 1990. Sprouting shrub response to different seasons and fuel consumption levels of prescribed fire in Sierra Nevada mixed co- nifer ecosystems. Forest Science 36:748—764. KoeEniIc, W. D., R. L. MumMmMeE, W. J. CARMEN, AND M. T. STANBACK. 1994. Acorn production by oaks in central coastal California: variation within and among years. Ecology 75:99-—109. McDOoNna_Lp, P. M. 1969. Silvical characteristics of Cali- fornia black oak (Quercus kelloggii Newb.). USDA Forest Service Research Paper PSW-53. . 1990. Quercus kelloggii Newb. California black oak. Pp. 661-671 in Silvics of North America, Vol. 2, hardwoods. USDA Forest Service Agricultural Handbook 654. AND D. W. Huser. 1995. California’s hardwood resource: managing for wildlife, water, pleasing scen- ery, and wood products. USDA Forest Service Gen- eral Technical Report GTR-PSW- 154. AND J. C. TAPPEINER. 1996. Silviculture-ecology of forest-zone hardwoods in the Sierra Nevada. Pp. 621—636 in Sierra Nevada Ecosystem Project: final report to Congress, Vol. III. University of California, Centers for Water and Wildlife Resources, Davis, CA. Multck, P. C. AND J. W. BARTOLOME. 1987. Factors asso- ciated with oak regeneration in California. Pp. 86—91 in Proceedings of the Symposium on Multiple Use of California’s Hardwood Resources. USDA Forest Ser- vice General Technical Report PSW-100. SAVAGE, M. 1994. Anthropogenic and natural disturbance and patterns of mortality in a mixed forest in Cali- fornia. Canadian Journal of Forest Research 24:1149— lis ep SPSS INCORPORATED. 1999. SYSTAT® 9 Statistics I. SPSS Incorporated, Chicago, IL. STANDIFORD, R. B., N. MCDOUGALD, W. FROST, AND R. PHILLIPS. 1997. Factors influencing the probability of oak regeneration on southern Sierra Nevada wood- lands in California. Madrono 44:170-183. , R. PHILLIPS, AND N. K. MCDOUGALD. 2000. The effect of thinning blue oak rangelands in California’s southern Sierra Nevada. Abstract 53rd Annual Meet- ing of the Society for Range Management, Boise, ID. ZAR, J. H. 1996. Biostatistical analysis, 3rd ed. Prentice Hall, Upper Saddle River, NJ. Maprono, Vol. 49, No. 2, pp. 122-129, 2002 CANOPY MACROLICHENS FROM FOUR FOREST STANDS IN THE SOUTHERN SIERRA MIXED CONIFER FORESTS OF SEQUOIA/KINGS CANYON NATIONAL PARK Davip C. SHAW Wind River Canopy Crane Research Facility, University of Washington, 1262 Hemlock Road, Carson, WA 98610 dshaw @u.washington.edu STEVEN A. ACKER Olympic National Park, 600 East Park Avenue, Port Angeles, WA 98362 ABSTRACT Canopy macrolichens were sampled using the “‘litter pickup” technique in four forest stands in the mixed conifer forests of Sequoia/Kings Canyon National Park. The purpose was to provide a basis for assessing lichen abundance trends in permanent forest plots, and to compare differences in lichen com- munities between four forest types typical of the southern Sierra Nevada. Each stand was characterized by a different conifer: sugar pine (Pinus lambertiana Dougl.), white fir (Abies concolor Gord. & Glend.), giant Sequoia (Sequoiadendron giganteum (Lindl.) Buchh.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.). The standing crop of lichen litterfall was estimated at 33.6 kg/ha, 14.8 kg/ha, 6.9 kg/ha, and 7.6 kg/ha respectively. Seven macrolichens were present in the litterfall, in decreasing order of overall abundance: Letharia vulpina (L.) Hue, Hypogymnia imshaugii Krog, L. columbiana (Nutt.) J. W. Thomson, Bryoria fremontii (Tuck.) Brodo & D. Hawksw. and Melanelia exasperatula (Nyl.) Essl., M. subolivacea (Ny1.) Essl., and Lobaria (Schreber) Hoffm. sp. A single factor ANOVA indicated that L. vulpina was equally distributed throughout the four stands, while H. imshaugii and L. columbiana were not. H. imshaugii was the most abundant lichen in the White Fir stand, although L. vulpina closely approximated it there. L. vulpina was most abundant in the Sugar Pine, Giant Sequoia and Jeffrey Pine stands, and all other lichens were much less abundant. A complex of factors explains the differences in lichen abundance; stand density, stand structure, and tree species composition appear most important, although site environmental conditions cannot be ruled out due to the lack of replication and small sample size in this study. The White Fir and Sugar Pine stands had 2—3 times the tree density as the Giant Sequoia and Jeffrey Pine stands. Giant sequoia and incense cedar (Calocedrus decurrens (Torr.) Florin) shed bark and therefore do not have abundant epiphytes on branches and tree boles. White fir appears to have a generally positive effect on lichen abundance, except in extremely dense stands. The abundance of H. imshaugii and L. columbiana were highly correlated with abundance of sugar pine. Although species diversity is low, standing crop of lichen litterfall is high, and may exceed many other forests in North America. Key Words: Macrolichens of forest canopies can be used to make inferences about a variety of ecosystem char- acteristics, including air quality, stand structure and history, stand age, and overall forest health (Segal and Nash 1983; Wetmore 1986; Boucher and Stone 1992; Bates and Farmer 1992; McCune 1993; Rhoades 1995). The distribution of these arboreal, non-crustose lichens across the landscape reflects the dynamic mosaic of environmental conditions (Hale 1974). Within the mixed-conifer forests of the southern Sierra Nevada Mountains in Califor- nia, canopy macrolichens have received limited study. The National Park Service and other government agencies are interested in determining whether the lichens are increasing or decreasing in abundance, because lichens may have value as indicators of environmental problems (McCune 2000). Smith (1980) did a taxonomic survey of the macrolichens in Sequoia/Kings Canyon National Park and found 40 species in 13 mostly forested study sites. An air Sierra Nevada, lichens, biomass, litterfall, canopy. pollution impact survey of all the lichens of Se- quoia/Kings Canyon National Park has identified 204 species (Wetmore 1986). Wetmore concluded that considering the dry climate, the lichen flora was diverse and healthy. The purpose of this study was twofold: 1. To document the relative abundance of canopy ma- crolichens in four forest stands that are part of a permanent forest plot system in Sequoia/Kings Canyon National Park (Harmon et al. 1987; Riegel et al. 1988). These data provide a baseline for fu- ture sampling to determine temporal trends in can- opy macrolichen abundance. 2. To compare the rel- ative abundance of canopy macrolichens in four forest stands dominated by different species of co- nifers and representing different environmental conditions in the lower montane, mixed conifer for- ests of the southern Sierra Nevada Mountains. The lower montane, mixed-conifer forests of the southern Sierra Nevada Mountains of California (1,600 m to 2,300 m) are characterized by giant 2002] sequoia (Sequoiadendron giganteum (Lindl.) Buchh.) (Cupressaceae), white fir (Abies concolor (Gord. & Glend.) Lindl. ex Hildebr.) (Pinaceae), California red fir (Abies magnifica A. Murr.) (Pin- aceae), sugar pine (Pinus lambertiana Dougl.) (Pin- aceae), Jeffrey pine (P. jeffreyi Grev. & Balf.) (Pin- aceae), and incense cedar (Calocedrus decurrens (Torr.) Florin) (Cupressaceae). Along a moisture gradient, giant sequoia occurs in mesic locations that do not dry out in the summer, white fir-mixed conifer (sugar pine and incense cedar) occurs in generally drier habitats, and Jeffrey pine occurs in the most xeric sites (Rundel et al. 1977; Vankat 1982). Fire and fire suppression play an extremely important role in stand composition and structure. In general, fire suppression results in an increase in the abundance of white fir (Rundel et al. 1977). Appropriate sampling for canopy macrolichen studies can be challenging, particularly for studies of trends in abundance over time. Canopy access using tree climbing is the most direct means of sampling canopy macrolichens, but sampling tree crowns to determine stand level abundance (i.e. biomass) requires very large amounts of time in tall forests (Clement and Shaw 1999). As an alterna- tive, McCune (1994) has developed a method to quantify the relative abundance of lichens in a for- est stand by sampling litterfall. This “‘litter pick- up’’ technique allows one to estimate the mass of canopy macrolichens at the stand level, which can then be used to compare relative abundance to other stand types and to determine stand-level trends in abundance over time. METHODS Study Site/Reference Stands The study site is located in the northwest portion of Sequoia National Park (Latitude 36°N and Lon- gitude 118°W) (Fig. 1). We chose four of the six reference stands described by Riegel et al. (1988), each dominated by a different species of conifer; Jeffrey pine, white fir, sugar pine (mixed conifer), and giant Sequoia (in a riparian setting). The stands are between 2,012 and 2,219 m in elevation and representative of three vegetation types: Sierran mixed-conifer (sugar pine and white fir), giant Se- quoia-mixed conifer (riparian), and Jeffrey pine (Riegel et al. 1988). The reference stands were es- tablished for long-term monitoring of vegetation and are cooperatively managed by the Sequoia/ Kings Canyon National Park, Oregon State Uni- versity, and the US Forest Service, Pacific North- west Research Station (Acker et al. 1998). The ref- erence stands were established in 1984 and re-mea- sured in June of 1994. All trees >5 cm have been tagged and mapped, and each tree has the diameter at breast height measured. Information is collected on crown ratio, crown vigor, tree mortality and damage. All data presented on stand structure comes from the 1994 measurement. SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON 128 Sequoia & Kings Canyon National GaN * FRESNO a A) A WOLVERTON Mixed Conifer SUWANEE CREEK HALSTEAD GENERAL SHERMAN TREE 124 > z + Riparian Giant a) yo Giant ) Sequoia CRESCENT MOW FOREST 4 MORO ROCK Fic. 1. Location of study site in Sequoia National Park, California (reproduced from Riegel et al. 1998). Four of these six reference stands were sampled, including White Fir, Jeffrey Pine, Riparian Giant Sequoia (Giant Sequoia) and Mixed Conifer (Sugar Pine). The study site has a mediterranean climate, with cool, moist winters and hot dry summers. Precipi- tation averages 1172 mm/year (1932—1983 mean at Giant Forest/Lodgepole, Sequoia National Park) and falls mostly as snow between November and April. The hot dry summers have a strong influence on arboreal lichen communities, which are charac- terized by low species numbers and dominance by a few drought-tolerant species. The Jeffrey Pine reference stand (1.0 ha) is on a moderately steep SE facing slope, (Table 1), with a glaciated granodiorite rock substrate. Exposed rock is common at the site. The canopy is open, domi- nated by Jeffrey pine (124 trees/ha), with California black oak (Quercus kellogii Newb.) (Fagaceae) (60 trees/ha) (Table 2). Dense clumps of shrubs, espe- cially green manzanita (Arctostaphylos patula Greene) (Ericaceae), are present. White fir (18 trees/ha), sugar pine (2 trees/ha) and incense cedar (5 trees/ha) occur in the lower plot where the slope flattens. This is a xeric, low productivity site. Jef- frey pine is adapted to the coarse textured soil found in the fissures of the glaciated granite (Riegel et al. 1988). The White Fir reference stand (0.9 ha) is located 124 MADRONO [Vol. 49 TABLE |. SITE CHARACTERISTICS OF THE FOUR REFERENCE STANDS IN SEQUOIA NATIONAL PARK. Reference Stand size Elevation Topographic Average stand (ha) (m) Aspect position slope (%) Sugar Pine 1.1 2091 southeast midslope-bench 11 White Fir 0.9 2012 southwest bench 20 Giant Sequoia 2.0 DANG) southwest/southeast lower slope 10 Jeffrey Pine 1.0 2109 southeast upper slope 23 on a flat area above the east-side of Suwanee Creek (Table 1). There are scattered outcrops of bedrock in the stand. The stand has a dense canopy of white fir (420 trees/ha) and California red fir (33 trees/ ha) near the stream which grades into a mixed-co- nifer forest with scattered sugar pine (56 trees/ha) and incense cedar (177 trees/ha) on the east side of the reference stand (Table 2). White fir is most abundant in all size- and canopy-classes,\while in- cense cedar, sugar pine and California red fir are more abundant in the intermediate and suppressed canopy classes and smaller diameter-classes (Riegel et al. 1988). The Sugar Pine reference stand (1.1 ha) is locat- ed to the west of Suwanee Creek approximately 200 m from the White Fir reference stand on a mid- slope bench (Table 1). The forest is a mosaic of large old sugar pine (110 trees/ha) and white fir (473 trees/ha) trees forming a relatively open can- opy in the dominant (sugar pine 20 trees/ha, white fir 19 trees/ha) and codominant (sugar pine 11 trees/ha, white fir 42 trees/ha) canopy classes (Table 2, Riegel et al. 1988). There are clumps of sup- pressed white fir and incense cedar (78 trees/ha) interspersed throughout the stand where white fir dominates the smaller size and canopy classes. Cal- ifornia black oak (16 trees/ha) and California red fir (7 trees/ha) are present in low numbers. The abundance of white fir in small size classes is thought to be a result of fire suppression (Riegel et al. 1988). The Giant Sequoia reference stand (2.0 ha) is on a lower slope, and straddles both sides of Crescent Creek (Table 1). There is a narrow corridor of her- baceous vegetation along the creek. The stand has a typical mixed conifer over-story dominated by gi- ant sequoia (24 trees/ha), which tower above the surrounding white fir (222 trees/ha) and California red fir (64 trees/ha) (Table 2). The true firs have a reverse J-shaped size distribution with a predomi- nance of small stems, as is typical of shade tolerant species (Riegel et al. 1988). Macrolichen Sampling Canopy macrolichens were sampled on June 20— 24, 1994 using 2-m radius (12.57 m7) litter pickup plots (McCune 1994). Litter refers to material Gin this case lichens) fallen from the canopy. At fifteen randomly chosen grid points in each reference stand, a stake was placed in the center of the plot and a 2-m string was used to measure the radius of TABLE 2. SPECIES COMPOSITION, NUMBER OF TREES PER HECTARE (TPH), TOTAL NUMBER OF SPECIES, DIAMETER (IN cm) CHARACTERISTICS, TREE SPECIES EVENNESS, TREE SPECIES RICHNESS BASED ON NUMBERS (KREBS 1989), TREE SPECIES RICHNESS BASED ON AREA (KREBS 1989) OF THE FOUR REFERENCE STANDS. Stand TPH by species Sugar Pine White Fir Giant Sequoia Jeffrey Pine White Fir 472.6 420.5 Dies 18.0 Red Fir Toll 33.0 63.5 0) Incense Cedar THD 177.3 0) 5.0 Jeffery Pine 0 0) 1.0 124.0 Sugar Pine 109.7 Se 6.0 2.0 Ponderosa Pine 0 0) 0) 1.0 Cal. Black Oak 15.9 0) 0 60.0 Giant Sequoia 0 0) D335) 0) TOTAL TPH 683.2 686.4 3525 210.0 # Tree Species 5) 4 5 6 median dbh Or 15.4 12.8 8.6 quad mean dbh 33.5 B72 81.3 31.8 max dbh 154.1 148.7 600.0 1333)5 I Basal area (m7?/ha) 60 74 164 17, Evenness 0.61 0.71 0.63 0.71 Rich No. 4.9 (0.27) 4.0 (0.00) 4.6 (0.51) 6.0 (0.00) Rich Area 5.0 (0.00) 4.0 (0.00) 4.7 (0.47) 5.9 (0.35) 2002] TABLE 3. TOTAL STANDING CROP OF CANOPY MACROLICHENS (kg/ha) ON THE FOREST FLOOR AND FREQUENCY OF OCc- CURRENCE IN 2 M RADIUS PLOTS. STANDARD DEVIATION IN PARENTHESES. SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON Stand Sugar Pine White Fir Giant Sequoia Jeffrey Pine Total Lichens Kg/ha 55:6 (27.7) 14.8 (18.5) 6:95:29) 7.6 (16.5) Frequency (%) 100 100 100 100 Letharia vulpina Kg/ha 15.04 (13.3) G5. (f233) 4.8 (1.5) T.33"(iG2) Frequency (%) 100 73 93 100 Letharia columbiana Kg/ha 4.4 (5.4) 095153) 0.2 (0.4) 0.15 (0.18) Frequency (%) 93 100 73 80 Hypogymnia imshaugii Kg/ha 14.1 (15.4) TS. (120) 8: (2:5) 0.08 (0.18) Frequency (%) 93 93 80 33 Bryoria fremontii Kg/ha 0.01 (0.03) 0) 0.08 (0.3) 0.01 (0.030 Frequency (%) 20 0 20 13 Melanelia spp. Kg/ha 0 0 0.06 (0.1) 0) Frequency (%) 0 0) 33 0 the plot. Flagging was located in four directions to denote the boundaries of the litter pickup plot. All fresh macrolichens (i.e., had not decayed beyond an identifiable state) were collected and placed in paper bags. Litter attached to wood was also col- lected, as was litter caught in shrubs up to 1 m off the ground that was not attached to the shrubs. Macrolichens were transported to the lab, cleaned, and sorted to species. The lichens were then dried at 60°C for 24 hours and weighed. Li- chen identifications were made using Hale and Cole (1988), and names follow Brodo et al. (2001). Spe- cies identifications were verified and unknown sam- ples were identified by Bruce McCune, Oregon State University. Reference specimens are deposit- ed in the University of Washington Herbarium. Analysis Biomass on each 2-m radius plot was trans- formed to g/ha for data analysis. The mean for the 15 plots in each reference stand was used to rep- resent stand level abundance and reported as kg/ha with standard deviation. Total lichen biomass and biomass of Letharia vulpina (L.) Hue (Parmeli- aceae), L. columbiana (Nutt.) J. W. Thomson (Par- meliaceae), and Hypogymnia imshaugii Krog (Par- meliaceae) were compared between reference stands using a single factor Analysis of Variance (Zar 1999), (d.f. = 3 between groups, and 56 d.f. within groups, a = 0.05). Although the study included only four stands, we explored various descriptors of forest structure and composition as potential predictors of total lichen litterfall biomass and biomass of each lichen spe- cies. Variation in lichen biomass was compared to variation in stand-level tree density and basal area, stem density of individual tree species, and tree species evenness and richness (from the rarefaction method (Krebs 1989)). RESULTS Species Seven species were found in the macrolichen lit- terfall of these four reference stands: Letharia col- umbiana, L. vulpina, Hypogymnia imshaugii, Bryoria fremontii (Tuck.) Brodo & Hawksw. (Par- meliaceae), Melanelia exasperatula (Nyl.) Essl. (Parmeliaceae), and M. subolivacea (Nyl.) Essl. (Parmeliaceae) and a Lobaria (Schreber) Hoffm. (Lobariaceae) sp. fragment. The Lobaria fragment was unidentifiable to species, and is not discussed further. The Letharia species and H. imshaugii were present in all four stands, while Bryoria fremontii was absent from the White Fir stand. The two Mel- anelia species were present only in the Giant Se- quoia stand which at six species, had the highest macrolichen litterfall species diversity. The other stands had four species, including the Lobaria sp. fragment at the White fir stand. Abundance The Sugar Pine stand had the highest standing crop of lichen litterfall (33.6 kg/ha) (Table 3). The White Fir stand had about % as much (14.8 kg/ha) and the Giant Sequoia (6.9 kg/ha) and Jeffrey Pine Biomass lichens (kg/ha) Number of Sugar Pine per hectare Fic. 2. Density of Sugar Pine per hectare on the four reference stands versus the standing crop of litterfall l- chen biomass per hectare for total lichens, Letharia vul- pina (LEVU), L. columbiana (LECO), and H. imshaugii (HYIM). Sugar Pine density corresponds to forest stands: = Jeffrey Pine, 6 = Giant Sequoia, 56 = White Fir, 110 = Sugar Pine. (7.6 kg/ha) stands had about one fourth that much lichen litterfall biomass as the Sugar Pine stand. In three of the four reference stands, lichen litterfall biomass was dominated by a combination of L. vul- pina and H. imshaugii. In the Giant Sequoia, White Fir, and Sugar Pine stands, L. vulpina accounted for 44% to 70% of lichen litterfall biomass and H. im- shaugii accounted for 25% to 49%. The Jeffrey Pine stand was unusual in that nearly all the lichen litterfall biomass (97%) was contributed by a single species, L. vulpina. The only other species to ac- count for 10% or more of the lichen litterfall bio- mass of any stand was L. columbiana, which was 13% of the biomass for the Sugar Pine stand. The ANOVA indicated significant differences between reference stands in biomass of total lichens (P < 0.01), L. columbiana (P < 0.01), and H. im- shaugii (P < 0.01). No significant difference was found for L. vulpina (P = 0.16). Biomass of L. columbiana and H. imshaugii generally increase with increasing density of sugar pine (Fig. 2). Tree density, basal area, tree species evenness, tree spe- cies richness based on numbers, or tree species richness based on area (Table 2, 3) shows little re- lationship to the variation in total biomass of li- chens. DISCUSSION Species Richness and Distribution The lichen litter pick-up technique documented only seven species of lichens in these four forest stands. This is low species diversity, even for dry habitats. Smith (1980) found 40 species of macroli- chens in the Ash Mountain to Grant Grove (High- way 198) region of Sequoia-Kings Canyon Nation- al Park and also included Cedar Grove. He sampled MADRONO [Vol. 49 13 sites using a floristic survey method that includ- ed all substrates, not just canopy lichens. The litter pick-up technique is not a ‘stand-alone’ method for surveys of species diversity, because species cap- ture is low. The technique is best used in conjunc- tion with other survey techniques that specifically search for different species of lichens (McCune and Lesica 1992; McCune 1994). However, it is a good technique for determining the relative abundance of the predominant forest canopy species. This is im- portant for monitoring change in lichen communi- ties. In a study of lichens on conifers and their rela- tion to air pollution in the Southern California mountains outside of Los Angeles, Sigal and Nash (1983) reported 16 species, the same number re- ported by Hasse for the same area in 1913 (Gin Sigal and Nash 1983). The lichen flora showed a strong relationship to air pollution: only eight of the orig- inal 16 species reported by Hasse were present in the most heavily polluted forests of the San Ber- nardino and San Gabriel Mountains. Sigal and Nash (1983) also rated the sensitivity of lichen species to air pollution, including several species we ob- served. They ranked L. vulpina as tolerant, M. su- bolivacea as moderately tolerant, and B. fremontii as very sensitive. Smith (1980) has given species accounts of all 40 species he observed in Sequoia/Kings Canyon National Park, including the six species observed in this study. According to Smith, Bryoria fremontii is uncommon, and was only collected once from the bark and branches of Pinus murrayana (Sierra lodgepole pine, P. contorta subsp. murrayana (Bal- four) Engelmann (Pinaceae)) in the Stony Creek area. We found B. fremontii in the Jeffrey Pine, Giant Sequoia, and Sugar Pine reference stands. Hypogymnia imshaugii was uncommon and was collected on A. concolor in the Crystal Cave Junc- tion area. Smith found H. enteromorpha (Ach.) Nyl. (Parmeliaceae) to be common and sometimes very abundant in all areas above 450 m. This in contrast to our finding of AH. imshaugii in all four sites, and a lack of collections for H. enteromorpha. Hale and Cole (1988) note that in the past, virtually all fertile Hypogymnias in California were called H.. enteromorpha, but that this name is now limited to populations along the coast that are characterized by grossly inflated branches, and that this species does not occur in the Sierra Nevada. Hale and Cole (1988) also indicate that H. imshaugii is very com- mon in Sequoia National Park. Smith considers Letharia columbiana and L. vul- pina to be two of the most common and abundant lichens in the park between 1200 m to 2700 m el- evation. He found them growing on numerous tree species all through the study region. We also found these two lichens to be abundant. Interestingly, L. vulpina was the more abundant of the two species with 3 to 10 times the biomass of L. columbiana in the reference stands. Smith found Melanelia subo- 2002] livacea (called Parmelia subolivacea Nyl.) abun- dant in all 13 study sites and in some trees the upper branches were completely covered by the li- chen. It was present on a wide variety of conifers and hardwoods. Parmelia exasperata De Not. was described as commonly found on Quercus, wide- spread in the Ash Mountain area, Potwisha, Buckey Flats and Deer Ridge. This may be what we iden- tified as M. exasperatula. We found these two spe- cies were present only in litterfall of the Giant Se- quoia reference stand. Abundance Letharia vulpina was the dominant lichen in three of these forest stands, and was generally equally distributed throughout the four forest stands. L. columbiana and H. imshaugii were not equally distributed and showed strong patterns of increase with increasing sugar pine and white fir. The extreme xeric conditions of the Jeffery Pine stand may have a negative influence on L. colum- biana and H. imshaugii. McCune (1994) has investigated canopy litter- fall relationships in the Pacific Northwest of North America. He found that the ratio 1:100 (litter : can- opy lichens) was fairly consistent in Douglas-fir forests for late summer standing crop of lichen lit- ter. Thus about 100 times the amount of lichen found on the forest floor in late summer will be in the canopy. This relationship has not been tested for forests of the Sierra Nevada. However if it is valid for the Sierra Nevada, the canopy biomass of macrolichens in the four reference stands would range from 0.7 Mg/ha in the Giant Sequoia stand to 3.4 Mg/ha in the Sugar Pine stand, with inter- mediate values for the Jeffrey Pine and White Fir stands (0.8 Mg/ha and 1.5 Mg/ha, respectively). These numbers are surprisingly large, perhaps in part because litter was collected in June rather than late summer. Typically a large pulse of lichen litter from winter storms will gradually disappear over the next 6-12 months depending on the spe- cies (McCune and Daley 1994). Another possibil- ity is that the mildly toxic Letharia spp are resis- tant to herbivory, resulting in greater persistence on the forest floor. Some of the most productive old-growth Doug- las-fir stands in the Pacific Northwest have 1.3 to 1.9 Mg/ha of macrolichens in the canopy (McCune 1993; McCune et al. 1997). Boucher and Nash (1990) estimated 0.75 Mg/ha macrolichens for can- opies of Blue Oak in California (36°N Latitude) while Turner and Singer (1976) estimated 1.9 Mg/ ha for a Pacific Silver Fir stand in the western Cas- cades of Washington. For further information on the biomass of epiphytes see Boucher and Stone (1992) and Rhoades (1995). The relationship of li- chen litter biomass to lichen biomass in the cano- pies of Sierra Nevadan forests is a key area for future research. SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON IDF Factors Influencing Lichen Abundance Lichen species composition and abundance in forest canopies is influenced by a multitude of fac- tors. Among other things, this includes tree spe- cies, bark texture/chemistry, stand age, ecological continuity of the forest (Bates and Farmer 1992), tree density, forest structure, disturbance history, air pollution, climatic conditions, and forest man- agement practices (Hale 1974; McCune 1993; Rhoades 1995). Within the southern Sierran mixed-conifer forests that we sampled, the most obvious influences on lichen species composition and abundance include tree species composition, stand density, and forest stand structure. It should be stressed that the generality of our interpretation is limited by the small sample size and no repli- cation of stand types. Differences in tree density did not directly cor- respond to differences in lichen biomass as stands with similar tree density differed in lichen biomass by a factor of 2 (Sugar Pine and White Fir) and stands with similar lichen biomass differed in tree density by 50% (Giant Sequoia and Jeffrey Pine). Tree species composition may explain some of these differences. Though they were similar in den- sity, the Sugar Pine and White Fir stands were very different in stand structure and species composition. The White Fir stand was uniform in stand structure creating more evenly shaded tree boles, and had Over twice aS many incense cedar (78/ha in the Sugar Pine to 177/ha in the White Fir stand). In- cense Cedar has exfoliating bark that sheds lichens. The Sugar Pine stand was more open with twice the number of sugar pine trees (110/ha in sugar pine to 56/ha in the fir stand) and a complement of dominant and codominant trees in the overstory, which provides for more sunlight on tree boles and branches. Thus, the Sugar Pine stand may have had an optimal combination of tree species composition and stand structure to provide for abundant lichen biomass. Giant Sequoia represented 74% of the basal area and 80% of the stand wood volume in the Giant Sequoia stand although accounting for only 7% of the stems. Giant sequoia also has exfoliating bark, hence the lichens are rare on the tree bole, and only abundant on dead wood and cones (Steve Sillett and Joel Clement, personal communication). This might explain why the Giant Sequoia and Jeffrey Pine stands were similar in lichen litterfall biomass even though the sequoia stand had 50% more trees. The Jeffrey Pine stand approached a woodland set- ting, with widely scattered trees, among outcrops of rock. Letharia vulpina was the dominant lichen in this forest, perhaps showing a tolerance for xeric conditions and compatibility for Jeffery pine bark texture and chemistry. The physical settings of the forest plots, such as aspect and proximity to streams, may also play a role in lichen abundance. The Giant Sequoia stand 128 MADRONO had a stream running through it, the White Fir stand and Sugar Pine stands were adjacent a stream, and the Jeffrey Pine stand was not influenced by a stream. A xeric to mesic environmental gradient was not measured in a systematic and replicated way in this study, and therefore conclusions re- garding the overall effect of tree species composi- tion as the major influencing factor associated with lichen abundance should be taken as a hypothesis needing further study. CONCLUSIONS We observed low species diversity of canopy h- chens in the mixed-conifer forests of the southern Sierra Nevada Mountains, yet an unusually high stand biomass of lichen litterfall. The Sugar Pine stand would be one of the highest biomass esti- mates for lichens in North America if the 1:100 ratio of litterfall to canopy lichen biomass for Northwestern forests (McCune 1994) holds true in Sierran forests. The early summer sample period and possibility of longer persistence on the forest floor by Letharia spp. may explain these higher numbers. Tree species composition (especially abundance of sugar pine), and canopy openness/ vertical structure appear to play a role in the abun- dance of canopy macrolichens, although the lack of replication within stand types and along the envi- ronmental moisture gradient preclude a definitive analysis. Characteristics of forest stands are con- trolled by a complex of factors, but in the future, anthropogenic influences such as fire suppression and controlled burning, air pollution, and climate change may become very important in determining lichen abundance. Long-term monitoring of lichens is important for understanding their role in the dy- namics of ecosystems and how they will respond to anthropogenic influences. ACKNOWLEDGMENTS This research was supported by the Wind River Canopy Crane Research Facility, a cooperative scientific endeavor of the University of Washington, Gifford Pinchot National Forest, and Pacific Northwest Research Station, US Forest Service, and the Sequoia/Kings Canyon National Park. Susan C. Shaw assisted with field sampling. Bruce McCune, Eric B. Peterson, Jeanne M. Ponzetti, and Tom Rambo provided editorial comment and review of the manuscript, which is especially appreciated. LITERATURE CITED ACKER, S. A., W. A. McCKEE, M. E. HARMON, AND J. FE FRANKLIN. 1998. Long-term research on forest dy- namics in the Pacific Northwest: a network of per- manent plots. Pp. 93-106 in F Dallmeier and J. A. Comiskey (eds.), Forest biodiversity in North, Central and South American, and the Caribbean: research and monitoring. UNESCO, Paris, France. BATES, J. W. AND A. M. FARMER (eds.). 1992. Bryophytes and lichens in a changing environment. Clarendon Press, Oxford, U.K. [Vol. 49 BOUCHER, V. L. AND T. H. NAsu, III. 1990. The role of the fruticose lichen Ramalina menziesii in the annual turnover of biomass and macronutrients in a blue oak woodland. Botanical Gazette (Chicago) 151:114—118. AND D. FE STONE. 1992. Epiphytic lichen biomass. Pp. 583-599 in G. C. Carroll and D. T. Wicklow (eds.), The fungal community: its organization and role in the ecosystem. Marcel Dekker, New York, NY. Bropo, I. M., S. D. SHARNOFF, AND S. SHARNOFF. 2001. Lichens of North America. Yale University Press, New Haven, CT. CLEMENT, J. P. AND D. C. SHAW. 1999. Crown structure and the distribution of epiphyte functional group bio- mass in old-growth Pseudotsuga menziesii tree crowns. EcoScience 6:243—254. HALE, M. E. Jr. 1974. The biology of lichens. Edward Arnold, London, U.K. AND M. Co_e. 1988. Lichens of California. Uni- versity of California Press, Berkeley, CA. HARMON, M. E., K. CROMACK, JR., AND B. G. SMITH. 1987. Coarse woody debris in mixed conifer forests in Se- quoia National Park. Canadian Journal of Forest Re- search 17:1265—1272. Kress, C. J. 1989. Ecological methodology. Harper and Row, New York, NY. McCune, B. 1993. Gradients in epiphyte biomass in three Pseudotsuga-Tsuga forests of different ages in west- ern Oregon and Washington. Bryologist 96:405—411. . 1994. Using epiphyte litter to estimate epiphyte biomass. Bryologist 97:396—401. . 2000. Lichens communities as indicators of forest health. Bryologist 103:353—356. AND P. LEsIcA. 1992. The trade-off between spe- cies capture and quantitative accuracy in ecological inventory of lichens and bryophytes in forests in Montana. Bryologist 95:296—304. AND W. J. DALy. 1994. Consumption and decom- position of lichen litter in a temperate coniferous rain- forest. Lichenologist 26:67—71. , K. A. AMSBERRY, FEF J. CAMACHO, S. CLERY, C. COLE, C. EMERSON, G. FELDER, P. FRENCH, D. GREENE, R. Harris, M. HUTTER, B. LARSON, M. LESKo, S. Ma- JORS, T. MARKWELL, G. G. PARKER, K. PENDERGRASS, E. B. PETERSON, E. T. PETERSON, J. PLATT, J. PROCTOR, T. RAmBo, A. Rosso, D. SHAW, R. TURNER, AND M. WIpMER. 1997. Vertical profile of epiphytes in a Pa- cific Northwest old-growth forest. Northwest Science 71:145-152. RHOADES, E M. 1995. Nonvascular epiphytes in forest canopies: worldwide distribution, abundance, and ecological roles. Pp. 353—408 in M. D. Lowman and N. M. Nadkarni (eds.), Forest canopies. Academic Press, San Diego, CA. RIEGEL, G. M., S. E. GREENE, M. E. HARMON, AND J. FE FRANKLIN. 1988. Characteristics of mixed conifer for- est reference stands at Sequoia National Park, Cali- fornia. Technical Report No. 32. Cooperative Nation- al Park Resources Studies Unit. University of Cali- fornia at Davis, Institute of Ecology, Davis, CA. RUNDEL, P. W., D. J. PARSONS, AND D. T. GORDON. 1977. Montane and subalpine vegetation of the Sierra Ne- vada and Cascade Ranges. Chapter 17 in M. G. Bar- bour and J. Major (eds.), Terrestrial Vegetation of California. John Wiley and Sons, New York, NY. SIGAL, L. L. AND T. H. Nasu III. 1983. Lichen commu- nities on conifers in southern California mountains: an ecological survey relative to oxidant air pollution. Ecology 64:1343—1354. 2002] SmiTH, D. W. 1980. A taxonomic survey of the macroli- chens of Sequoia and Kings Canyon National Parks. M.S. Thesis. San Francisco State University, San Francisco, CA. TURNER, J. AND M. J. SINGER. 1976. Nutrient distribution and cycling in a sub-alpine coniferous forest ecosys- tem. Journal of Applied Ecology 13:295-—301. VANKAT, J. L. 1982. A gradient perspective on the vege- SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON 129 tation of Sequoia National Park, California. Madrofio 29:200-214. WETMORE, C. 1986. Lichens and air quality in Sequoia National Park and Kings Canyon National Park. Sup- plementary Report, Contract CX 001-2-0034. Nation- al Park Service, Denver, CO. ZAR, J. H. 1999. Biostatistical analysis. Prentice-Hall, En- glewood Cliffs, NJ. MApRONO, Vol. 49, No. 2, pp. 130-131, 2002 CALYSTEGIA SILVATICA (CONVOLVULACEAE) IN WESTERN NORTH AMERICA R. K. BRUMMITT The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K. At the kind request of the Jepson Manual editor, I contributed the account of Calystegia to the recent edition of the Jepson Manual (Brummitt in Hick- man 1993: 517-521) despite my never having set foot in California at the time. My knowledge of the taxa in California, where half the genus is endemic, was based entirely on extensive study of herbarium specimens sent on loan from the major herbaria in the late 1960s. I was unaware of any introduced species established in the state, though I was fa- miliar with several species of the genus which had become very well naturalized as weeds in various other parts of the world. In 1997 I attended a meeting at the California Academy of Sciences, and, to my great delight, was able to see in the field for the first time many of the Calystegia taxa native to the state. But, to my surprise, I was able to add one introduced species to the Californian list before ever stepping into the field, when I reidentified a specimen in the CAS collections as C. silvatica (Kit.) Griseb., native of southern Europe. The specimen was collected at Stinson Beach in Marin Co. in the late 1950s by J.T. Howell and then identified and laid away as C. sepium (L.) R.Br. My host at CAS, Tom Daniel, immediately agreed to take me to Stinson Beach to see if the plant was still there, and, sure enough, it was there in great abundance on waste ground over a distance of perhaps 100 m, swamping other veg- etation and climbing to a height of 3 m, very con- spicuous even to a passing motorist. It was surpris- ing that such a conspicuous plant has been so over- looked, and we collected additional material: Marin Co., Stinson Beach, roadside in town, alt. 20 m, 122°39’W, 37°54'N, abundant, rampant over vege- tation to 3 m, 31 May 1997, R.K. Brummitt & T.F. Daniel 19614 (CAS, K, MO, RSA, UC). Alerted to the presence of this species in the area, I noticed it again a few days later some 11 km east of Stinson Beach on waste ground in the town of Mill Valley, and collected it again: Marin Co., Mill Valley, near foot of Reed Street, roadside in sub- urban area, 122°33'W, 37°54'N, climbing over bushes and roadside vegetation to 3 m, 10 June 1997, R.K. Brummitt 19672 (CAS, K). Again, the strongly climbing stems and the large white flowers were very conspicuous. C. silvatica is readily distinguished from any Calystegia in northern or mid California by its very vigorous habit, large glossy leaves, and particulaly its large flowers (corolla 50-75 mm) with two large braceoles at the base which are inflated and over- lapping and more or less obscuring the calyx. In southern California it is approached in its flowers and bracteoles by C. macrostegia (House) Brum- mitt subsp. macrostegia and subsp. amplissima Brummitt from the Channel Islands, but that spe- cies is woody at the base whereas C. silvatica is rhizomatous, and the two species are almost cer- tainly not closely related despite their superficial similarity. The pan-temperate complex of C. sepium and re- lated species, including C. silvatica, is difficult to resolve taxonomically, consisting of a series of geo- graphical taxa with minor distinguishing characters. My concept of C. silvatica includes plants native of eastern N. America, southern Europe and as far east as Iran, and China, which occur sympatrically with C. sepium except in China and are distin- guished from it by their large inflated and overlap- ping bracteoles. Those from eastern N. America and China are characterised by sometimes having twin peduncles in leaf axils and a rather square leaf sinus, and I refer them to subsp. fraterniflora (Mack. & Bush) Brummitt. In the Mediterranean region those from central and southern Italy east- wards have a longer range of flower size and big- ger, more inflated bracteoles with a rounded to emarginate apex, and these are referred to subsp. silvatica. Plants from northern Italy and southern Switzerland westward to Spain tend to have a smaller range of flower size and an obtuse apex to the bracteoles. Despite more than thirty years of hesitation over whether to give this variation formal taxonomic recognition—which necessitated a lengthy note on the matter in Flora Europaea (Brummitt 1972) instead—I have recently formally separated these southwestern European plants as subsp. disjuncta Brummitt (see below). Both of the Mediterranean subspecies were in- troduced into the British Isles about a century ago, probably as garden ornamentals, and both are now serious weeds there (but, curiously, not so in other north European countries). Subsp. silvatica is also known as a weed in Australia, where subsp. dis- juncta appears not to have been introduced. In North America, however, it seems to have been only subsp. disjuncta that has become established, this being known from specimens collected in Washington State from 1927 onwards and also from British Columbia and Oregon. Its occurrence in California is thus not very surprising. Subsp. fra- 2002] terniflora has been recorded as a rare alien in the British Isles, almost certainly introduced from North America, but has not become established. In the standard text on the plants of the Pacific Northwest, Hitchcock (1959) included all these taxa in Convolvulus sepium L. The plant figured there under this name on p. 88 is very probably C. sil- vatica subsp. disjuncta. The plant referred to in the text as var. fraterniflorus is probably also this taxon and this epithet is misapplied here. The statement that it is native of the eastern United States is in- correct. It was already described by Hitchcock here as a difficult weed. All taxa in the C. sepium complex spread vig- orously by rhizomes and tend to become aggressive weeds, often swamping the vegetation the stems climb over. The rhizomes are, however, not quite so deep-rooted as those of Convolvulus arvensis L., a major weed in California, and they are not quite such a persistent pest. Seed dispersal in C. sepium and allies is less significant than vegetative spread, since there is no obvious dispersal mechanism and the seeds tend to merely fall to earth beneath the parent plant. Furthermore, all plants in this complex are self-sterile and single populations normally do not set seed at all. C. silvatica tends to be more vigorous in its vegetative spread than the variants of C. sepium are, and its potential as a serious weed is considerable. It seems that in Britain new popu- lations may be established by accidental transport of pieces of rhizome (see, for example, notes in Brummitt & Chater 2000). Calystegia silvatica (Kit.) Griseb., Spic. Fl. Rum. 2: 74 (1844) subsp. disjuncta Brummitt in La- gascalia 18: 339 (1996). Additional specimens documenting known range in North America. BRITISH COLUMBIA. West Vancouver, 16 Ma- BRUMMITT: Calystegia silvatica in Western North America 131 rine Drive, roadside, 6 July 1955, W. Bird 1239 (BM); Vancouver, Bush Crown & 25 Ave., 5 May, 1956, W. Bird 2008 (BM): Vancouver, Canoe Pass Delta, bank between road and river, 1 Sept. 1957, W. Bird 3533 (BM). WASHINGTON. Marysville, fields, July 1927, J.M. Grant (US); Bank of R. Spokane, July 1931, Sister M. Milburge (WTU); Seattle, end of Green Lake, dumping ground, 28 July 1933, J.W. Thomp- son 9619 (NY); Port Blakely, in thicket, 17 June 1934, W.J. Eyerdam (L, MO); Whatcom Co., Birch Bay, 19 July 1937, W.C. Muenscher 8343 (DAO); King Co., Fort Lawton, roadside hedges, 25 July 1937, W.J. Eyerdam (EK MO); % mile NE of Seattle, near Sand Point, in thicket, 10 June 1949, W.J. Eyerdam (BM); Shelton—Woodsport road, moist sandy soil, 30 July 1950, P.E. Freer 320 (WTU). OREGON. Just east of Corvallis, across Van Bu- ren St. bridge, 15 June 1960, L.R.J. Dennis 2227 & G. van Vechten (NY). It is a pleasure to thank many botanists we have assisted me in the field in California on my three visits to look at Calystegia in 1997, 1998 and 1999, especially Barbara Ertter (Berkeley) and Tom Dan- iel (California Academy of Sciences). LITERATURE CITED BRUMMITT, R. K. 1972. Calystegia. Pp. 78-79 in T. G. Tutin et al., (eds.), Flora Europaea, Vol. 3. Cambridge University Press, Cambridge, U.K. BRUMMITT, R. K. AND A. O. CHATER. 2000. Calystegia (Convolvulaceae) hybrids in West Wales. Watsonia 23:161—165. HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. Hitcucock, C. L. 1959. Pp. 85-89 in C. L. Hitchcock, A. Cronquist, and M. Ownbey (eds.), Vascular Plants of the Pacific Northwest, Vol 4. University of Washing- ton Publications in Botany, 17. University of Wash- ington Press, Seattle, WA. MADRONO, Vol. 49, No. 2, pp. 132—133, 2002 NOTEWORTHY COLLECTIONS ARIZONA MANCOA PUBENS (A. Gray) Rollins (BRASSICA- CEAE).—Cochise County, San Pedro Riparian National Conservation Area, Kolbe site ca. 2 km SW of San Pedro Inn Bed and Breakfast, ca. 3 km S of Hereford Bridge, ca. 100 m E of San Pedro River. N31°24.961' W110°6.227’, elevation 1274 m, June 6, 2001. Found in sacaton grassland habitat with young mesquite; upper flood plain in sandy-loamy soil; level, open aspect. As- sociated species include Sporobolus wrightii, Prosopis ve- lutina, Erigeron concinnus, Xanthocephalum gymnosper- moides, Helenium thurberi, Conyza coulteri, Conyza can- adensis, Plantago sp., Salix goodingii, Elizabeth Makings 365 (ASU). Previous knowledge. West Texas to Coahuila, Mexico (Rollins, The Cruciferae of Continental North America, Stanford University Press, 1993). Other regional collec- tions: Hinckley s.n. (ARIZ) Jefferson Davis County, Texas in 1937; Waterfall s.n. (ARIZ) Jefferson Davis County, Texas in 1943; Andres Rodriguez 861 (TEX) Coahuila, Mexico in 1983; four collections from Jefferson Davis County, Texas (TEX); two collections from Presidio County, Texas (TEX); and four collections from Brewster County, Texas (TEX). Significance. First record for the species in Arizona. This collection is approximately 500 km from nearest known collections in Jefferson Davis County, Texas. This was an isolated annual/biennial, inconspicuous in its in sacaton grassland community, which may explain why it has been under collected. This habitat is widely recog- nized as a major corridor for migratory birds. Seed size make wind dispersal unlikely, therefore, bird dispersal may explain the considerable range extension for this spe- cies. —ELIZABETH MAKINGS, Department of Plant Biology, Arizona State University, PO. Box 871601, Tempe, AZ 85287-1601. W ASHINGTON BACCHARIS PILULARIS DC (ASTERACEAE).—Pacific Co: Fort Canby State Park, Beard’s Hollow. R11W, TION, S23, SW % of NE %4, USGS 7.5 minute “Cape Disap- pointment”’ quad, 30 m SW of Beard’s Hollow overlook on State Route 100, 1.5 km W of Ilwaco. Growing on SW facing cliffs in low coastal seacliff meadows above dune forest, with Poa unilateralis, Festuca rubra, Sedum ore- gonensis, Vulpia bromoides, and below Calamagrostis nutkaensis meadows. Colony of 30-50 upright plants, 0.5—2 m tall, at 10-30 m elevation. One plant grows on sand 100 m to the north, at an elevation of 4 m, with Alnus rubra, Ammophila arenaria, Leymus mollis. First seen in 1981, revisited and collected while in flower on 9-15-2001. K. Sayce (WS, OSC, WTU). Previous knowledge. Baccharis pilularis occurs from Oregon to northern Mexico on coastal bluffs to oak wood- lands, occasionally on serpentine, 0-750 (1500) m. The nearest known occurences are Gearhart and Cannon Beach, Clatsop County, Oregon (information supplied by Richard Halse, OSU Herbarium, Corvallis). Baccharis pi- lularis is increasingly common to the south, and is a mem- ber of shrub communities, including chaparral and coastal sage, in California. Significance. This is the northernmost known location and only Washington site for Baccharis pilularis, and is 30 km north of the next known site in Oregon. This small population may represent remnants of a species that was more widespread in warmer climates. —KATHLEEN SAYCE, P.O. Box 91, Nahcotta WA 98637. kas @sbpac.com Moorea, SOCIETY ISLANDS, FRENCH POLYNESIA ARUNDO DONAX L. (POACEAE).—Poa Poa, University of California, Gump Biological Research Station, hillside grounds of main house and beyond, 17°30’S, 149°49’'w, elev. 100 m, 25 October 2000, Mitchel P. McClaran and James W. Bartolome 00-04 (ARIZ, UC). Previous knowledge. Native to Mediterranean (Allred 1993, In: Hickman (ed.), The Jepson Manual, University of California Press, Berkeley, CA, p. 1235) cultivated worldwide, and regularly naturalized. Known from nearby Huahine and Raiatea islands (Welsh 1998, Flora Societen- sis, Electronic PrePrint Services Inc., Orem, UT). Significance. First record of this non-native species for Moorea. Several large plants spreading from landscape grounds to wild hillside. Based on evidence from Southern California, this species spreads via vegetative reproduc- tion, can dominate riparian vegetation through competitive exclusion, and can facilitate an increase in fire occurrence (Bell 1997, In: Brock et al. (eds.). Plant Invasions: Studies from North America and Europe. Backhuys Publishers, Leidens, Netherlands, p. 103-113). CHLORIS BARBATA Sw. (POACEAE).—Poa Poa, in waste area surrounding Supermarche Aré market, 17°29’S, 149°49’'W, elev. 2 m, 23 October 2000, Mitchel P. Mc- Claran and James W. Bartolome OO-O1 (ARIZ, BISH, UC). Previous knowledge. Native to New World tropics and naturalized worldwide, and known from nearby Tahiti and Raiatea islands (Welsh 1998). Significance. First record of this non-native species for Moorea. Naturalized and common along roadsides and other waste areas near human settlements. This distribu- tion 1S consistent with observations from other South Pa- cific archipelagoes, including Hawaii, Samoa, Tonga, Fiji, Guam, and Belau (Whistler 1995, Wayside Plants of the Islands, Isle Botanica, Honolulu, HI). HYPARRHENIA RUFA (Nees) Stapf (POACEAE).—ca. 2 km S of Afariatu, near and in abandoned livestock pasture, assoc. with Setaria sphacelata (Schumach.) Stapf & C.E. Hubb. ex. M.B. Moss, 17°34’S, 149°47'W, elev. 4 m, 24 October 2000, Mitchel P. McClaran and James W. Bar- tolome 00-02 (ARIZ, BISH, UC). Previous knowledge. Native to tropical Africa and America (Renvoize 1984, The Grasses of Bahia, Kew Bo- tanical Garden, England). Known from nearby Tahiti and Raiatea islands (Welsh 1998). 2002] Significance. First record of this non-native species for Moorea. Not common, but plants present beyond bound- aries of area seeded for livestock pasture. Naturalized, and likely to continue to spread based on experience in Aus- tralia, Hawaii, and Venezuela. Considered a weed in Aus- tralia because it spreads from seeded areas into native veg- etation (Lonsdale 1994, Australian Journal of Ecology 19: 345-354). Spread increases the occurrence of fire, which hastens its increase and the decrease of native species in Hawaii and Venezuela (Smith and Tunison 1992, In: Stone et al. (eds.), Alien Plant Invasions in Native Ecosystems of Hawaii: Management and Research, University of Ha- wail Cooperative National Park Resources Study Unit, Honolulu, HI, p. 394—408, and Baruch 1996, In: Solbrig et al. (eds.), Biodiversity and Savanna Ecosystem Pro- cesses: A Global Perspective, Springer-Verlag, New York, NY, p. 79-93), although it will spread and dominate native vegetation in the absence of fire and grazing (San Jose and Farinas 1991, Acta Oecologia 12:237—247). SETARIA SPHACELATA (Schumach.) Stapf & C.E. Hubb. ex. M.B. Moss (POACEAE).—ca. 2 km §S of Afariatu, near and in abandoned livestock pasture, assoc. with Hy- parrhenia rufa (Nees) Stapf, 17°34'S, 149°47'W, elev. 4 m, 24 October 2000, Mitchel P. McClaran and James W. Bartolome 00-03 (ARIZ, BISH, UC). NOTEWORTHY COLLECTIONS 133 Previous knowledge. Native to tropical and subtropical Africa, and Yemen (Hacker 1992, In: ‘t Mannetje & Jones (eds.), Plant Resources of South-east Asia, No. 4, Pudoc Sci. Publ., Wageningen, Netherlands, p. 201—203). Known from seeded pasture, Taiarapu Plateau on nearby Tahiti island (Welsh 1998). Significance. First record of this non-native species for Moorea. Not common, but plants present beyond bound- aries of area seeded for livestock pasture. Possibly less likely to spread from seeded area than Hyparrhenia rufa based on its absence from Australian weed lists (Lonsdale 1994). However, that prediction may be incorrect because this collection is apparently the more fecund cultivar ‘Splenda’, based on intermediate values for some char- acters (number of culm nodes, culm diam., lf width, and panicle length) between vars. sericea and _ splendida (Hacker 1992). This cultivar, developed in Australia in the 1980s, had high yields in Southeast Asian and Southern Pacific trials, and unlike the two varieties, will produce fertile seeds in abundance (Hacker 1992). —MITCHEL P. MCCLARAN, School of Renewable Natural Resources, 325 Bioscience East, University of Arizona, Tucson, AZ 85721 and James W. Bartolome, Department of Environmental Science, Policy, and Management, 145 Mulford Hall, University of California, Berkeley, CA 94720. MADRONO, Vol. 49, No. 2, p. 134, 2002 REVIEW Seeing things whole: the essential John Wesley Powell. Edited by WILLIAM DEBuys. 2001. Island Press, Covelo, CA. 388 pp. $27.95. ISBN 1-55963- 872-9. Guessing right about the direction history will take is always a doubtful proposition, but as Wil- liam deBuys makes clear in Seeing things whole: the essential John Wesley Powell, the famous one- armed explorer and scientist did guess right about enough important issues of his time to still be worth reading today. Actually, as deBuys makes clear, to say that Powell “‘guessed right’’ is not to do him the justice he deserves. A true polymath, Powell spent decades studying the landscape and peoples of the region he cared about—the American West. First as ex- plorer of the Grant Canyon, and then, over time, as geologist, ethnologist, bureau administrator, social planner, public figure, and philosopher, Powell worked over the same territory again and again from endlessly evolving perspectives, seeing pat- terns where his contemporaries clearly did not. Powell has not been ignored by the historians of recent times, and it is a tribute to the Major’s sig- nificance that he has been the subject of biographies by both Wallace Stegner and Donald Worster. What deBuys adds to these two excellent histories is Powell in his own words. This selection of Powell’s writings varies from excerpts from his well-known Colorado River writings, to things much less known including such forgotten statements as the Major’s address to Montana’s 1889 constitutional convention. Particularly welcome are several chap- ters from Powell’s often quoted (but not so often read) Report on the arid lands. What deBuys shows us is not a Powell who es- caped all the constraints of his times, but rather a powerful and original thinker who worked hard to see what was going on in his time. Certainly, by contemporary standards, Powell could not escape 19th century expectations that the native people of North America were destined to disappear, and as an ecologist he had trouble (as many still do) com- ing to grips with the role of the fire in the West, but what deBuys shows clearly is that in a number of critical areas Powell did break through into new ways of thinking about a number of still critical issues. Certainly, his recognition that the nation’s land laws would victimize both many of those who tried to settle the West and the land itself, was un- precedented and still highly valid today. William deBuys is to be thanked for bringing a useful se- lection of Powell’s original writings back into easy reach. —WILLIAM TWEED. Sequoia-Kings Canyon National Parks, Three Rivers, CA 93271. Maprono, Vol. 49, No. 2, p. 135, 2002 REVIEW Inventory of rare and endangered plants of Cali- fornia, sixth edition. California Native Plant Soci- ety, Rare Plant Scientific Advisory Committee, Da- vid P. Tibor, convening editor. 2001. California Na- tive Plant Society, Sacramento, CA. 388 pp. Soft- cover $29.95. ISBN 0-943460-40-9. The California Native Plant Society’s recent pub- lication of the sixth edition of the Inventory of rare and endangered plants of California offers a wel- come update on the constantly changing condition of endangered plants and habitats. Plants species’ rare, threatened or endangered status has expanded by a hefty 19% since the fifth edition, issued in 1994, so it is an essential addition to any library. This version basically adheres to the same suc- cessful formula of its predecessors. However, some new features that appear in this /nventory are the inclusion of rare, non-vascular bryophytes (mosses, liverworts and hornworts), each species’ typical el- evation ranges, county indexing for all 2073 plants, and a new common name appendix. Inside covers have helpful lists of often-used, and too-often for- gotten acronyms. The Jnventory is a massive collaborative effort utilizing the talents of hundreds of scientific con- tributors who have spent untold hours in the field identifying, mapping, and gathering botanical data to be added to or updated in the Department of Fish and Game database. The importance of this work has its basis in that critical and balanced scientific input. But with a readership as diverse as devel- opers, biological consultants, educators and private landowners, the /nventory also must and does adeptly make this information understandable and available to all intellectual levels, thereby providing the greatest opportunity for early detection as well as avoidance of potential environmental conflicts. Introductory chapters espouse both CNPS poli- cies as well as those of various state & federal agencies—simplifying and clarifying the some- times overwhelming mass of plant protection guidelines and legislations. Peggy Fiedler again provides the scientific rationale for the plant endan- germent rating system. The actual “Threatened and Endangered Plant In- ventory’ is alphabetically organized by scientific name for easy access. In most cases, specific entries contain the following information: scientific names as well as family and common names, associated authorities, CNPS’ rarity rating (and those given state and Federal status as well), occurrences by county with map coordinates, key identifying char- acteristics, typical habitat and pertinent historical notations with literary referencing. The Inventory of rare and endangered plants is one of the most tangible and fruitful of CNPS’ many pursuits. Convening Editor Tibor and his le- gions of collaborators have not only achieved their intent for the /Jnventory, which encourages “ conservation planning and enforcement of environ- mental laws that protect rare species,’ they have ably furthered the goals of the California Native Plant Society. —MELANIE BAER-KEELEY. Division of Resources Man- agement, Branch of Vegetation Management, Sequoia Kings Canyon National Park, Three Rivers, CA 93271. MADRONO, Vol. 49, No. 2, p. 136, 2002 CALIFORNIA BOTANICAL SOCIETY 2002—2003 SCHEDULE OF SPEAKERS All Meetings are held at 7:30 p.m. on the 3rd Thursday of the month Sepia Oct. 17 Nov. 21 Jan. 16 Feb. 15 Mar. 20 April 17 May 15 Room 2063, Valley Life Sciences Building University of California, Berkeley Martin Bidartondo, Dept. of Plant Biology, UC Berkeley Obligate cheaters of mycorrhizal networks. Truman Young, Dept. of Environmental Horticulture, UC Davis Ecological restoration: an emerging conservation strategy. Randy Jackson, ESPM, UC Berkeley Spring-fed wetland structure and function in California oak savannas. David Ackerly, Dept. of Biological Sciences, Stanford University Fire, drought, and the evolution of chaparral shrubs. Annual banquet (University of San Diego) Jon Rebman, San Diego Natural History Museum Discoveries on a floristic frontier: Baja California Kim Steiner, California Academy of Sciences The evolution of a specialized pollinator system in southern Africa. Todd Dawson, Dept. of Integrative Biology, UC Berkeley Giants in the mist: coastal redwoods and the land-sea interface. Marcel Rejmanek, Div. of Biological Sciences, UC Davis Seed dispersal and coexistence of tree species in tropical forests Open to non-members. Refreshments served after the presentation. Volume 49, Number 2, pages 61—136, published 17 December 2002. —————<«— —. SUBSCRIPTIONS—MEMBERSHIP Membership in the California Botanical Society is open to individuals ($27 per year; family $30 per year; emeritus $17 per year; students $17 per year for a maximum of 7 years). Late fees may be assessed. Members of the Society receive Maprono free. Institutional subscriptions to MADRONO are available ($60). Membership is based on a calendar year only. Life memberships are $540. Applications for membership (including dues), orders for sub- scriptions, and renewal payments should be sent to the Treasurer. 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Printer’s fees for illustrations and typographically difficult material @ $35 per page (if their sum exceeds 30 percent of the paper) and for author’s changes after typesetting @ $4.50 per line will be charged to authors. At the time of submission, authors must provide information describing the extent to which data in the manu- script have been used in other papers that are published, in press, submitted, or soon to be submitted elsewhere. VOLUME 49, NUMBER 3 JULY-SEPTEMBER 2002 RECEIVED BY: §§ F / INDEXING (-/- IND RIAN PENINSULA AND THE BALEARIC ISLANDS Roberto Gamarra.and Cindy Talbott ROCHE s.:....0020002000cs0eecsneceacessaseneenoens 7 HyYMONOCLEAS ARE AMBROSIAS (COMPOSITAE) Jonni Strother and Bruce G. BOVAWUD so.diccescsssesscastasaincewaeedcaconaseesvaraeenes 143 THE FLORA OF ToRA ISLET AND NOTES ON GUADALUPE ISLAND, BAJA CALIFORNIA, MEeExIco Jon P. Rebman, Thomas A. Oberbauer, and José Luis Leén de la Luz.... 145 GENETIC STRUCTURE OF SENECIO LAYNEAE (COMPOSITAE): A RARE PLANT OF THE CHAPARRAL Glenda D. Marsh and Debra R. AYTES. «.polgg¥2)...2<02000:20pPh- aisle nonacsesesesestes 150 Woop ANATOMY AND SUCCESSIVE CAMBIA IN SIMMONDSIA (SIMMONDSIACEAE): EVIDENCE FOR INCLUSION IN CARYOPHYLLALES S. L. SHOR WOR COI GUISE cook se mcncecc Ath Bc acacin aac ee NOR << pea cc. Notes svesencnetacescnes 158 A NEw SUBSPECIES OF NAVARRETIA LEUCOCEPHALA (POLEMONIACEAE) FROM VERNAL POoLs IN EASTERN WASHINGTON Curtis R. BjOrRE Serge AOR A ee ON at oo RI). nn ceg cess ceeees 165 RESURRECTION OF A CENTURY-OLD SPECIES DISTINCTION IN CALAMAGROSTIS Barbara YWAIsOn GAG SGHIMGFOVR Sr Nase IER wating nnn ee ToT OAR cones 169 PLANT ZONATION IN A SHASTA COUNTY SALT SPRING SUPPORTING THE ONLY KNOWN POPULATION OF PUCCINELLIA HOWELLII (POACEAE) Lary evng Mary Bat ca GAD KAO¥ ul CRA re srs ikon eee cecceceeeseee 178 SPHAGNUM BALTICUM IN A SOUTHERN RocKY MOUNTAIN IRON FEN David J. Cooper, Richard E. Andrus and Christopher D. Arp ..............+++- 186 EVIDENCE OF A NOVEL LINEAGE WITHIN THE PONDEROSAE AnniM. Patten and Steven I. Bi HgfOl aed oo..c.ccscoincscSMede ah on odeevecasecosseseesse 189 JTURSTOR pool acca ey Ueno Se | 22 eee 193 COULTER 22 28 Re SAO oe 2 a See ae ena oe eee 193 Ones re nee a ne See oe he eae «lj cusp vu Vana encvarsiuwassnendetcnveadecs 194 oy a STEPTGHIGIN), 2s. Sees 8 00h 09 of Spa) AO Er atene On fee) Se ee ee ee ee 195 A Cactus OpyssEy: JOURNEYS IN THE WILDS OF BOLIVIA, ARGENTINA, AND PERU, BY JAMES D. MAuUSETH, ROBERTO KIESLING, AND CARLOS OSTOLAZA Mae cateeBtatea Cork FAG NN TE ee ea note ens a er Setice tobi o css onaces Suansdinesedsibdieibineueseecres 198 BIENNIAL GRADUATE STUDENT MEETING AND ANNUAL BANQUET ..........cesceecceecceeceeees 199 Eon A eae ys re hoe oan Raney sive cddeesevaserassaswcevesene 200 Maprono (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. PostmMasTER: Send address changes to MADRONo, Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. Editor—Dnr. JOHN CALLAWAY Dept. of Environmental Science University of San Francisco 2130 Fulton Street San Francisco, CA 94117-1080 callaway @usfca.edu Book Editor—Jon E. 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Sciences, One Washington Square, San José, CA 95192. rmyatt@email.sjsu.edu Second Vice President: MicHAEL S. Mayer, Department of Biology, University of San Diego, San Diego, CA 92110, mayer @sandiego.edu Recording Secretary: | Stact Markos, Friends of the Jepson Herbarium, University of California, Berkeley, CA 94720- 2465, smarkos @socrates.berkeley.edu. Corresponding Secretary: | SUSAN BAINBRIDGE, Jepson Herbarium, 1001 VLSB #2465, University of California, Berkeley, CA 94720-2465. 510/643-7008. suebain@SSCL.berkeley.edu Treasurer: Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. The Council of the California Botanical Society comprises the officers listed above plus the immediate Past President, R. JouN LittLe, Sycamore Environmental Consultants, 6355 Riverside Blvd., Suite C, Sacramento, CA 95831; the Editor of Maprono; three elected Council Members: JAMES SHEVOCK, National Park Service, 1111 Jackson St., Suite 700, Oakland, CA 94607-4807. 510/817-1231; ANNE BRADLEY, USDA Forest Service, Pacific Southwest Region, 1323 Club Drive, Vallejo, CA 94592. abradley @fs.fed.us; DEAN KELCH, Jepson and University Herbarium, University of California, Berkeley, CA 94720, dkelch @sscl.berkeley.edu. Graduate Student Representative: ELIZABETH ZACHARIAS, Department of Integrative Biology, University of California, Berkeley, CA 94720. ezachar@socrates.berkeley.edu; Local (San Diego) Graduate Student Representative: RoBErT K. Lauri, Department of Biology, San Diego State Uni- versity, San Diego, CA 92182. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Maprono, Vol. 49, No. 3, pp. 137—142, 2002 DISTRIBUTION OF THE GENUS CRUPINA IN THE IBERIAN PENINSULA AND THE BALEARIC ISLANDS ROBERTO GAMARRA Dpto. de Biologia (Botanica), Facultad de Ciencias, Universidad Autonoma de Madrid, Cantoblanco, E-28049 Madrid, Spain CINDY TALBOTT ROCHE* 109 Meadow View Drive, Medford, OR 97504 USA crupinaqueen @charter.net ABSTRACT Crupina vulgaris, with populations in Sonoma and Modoc counties, is a relatively recent Mediterranean invader among the 1045 alien plant species naturalized in California, apparently introduced from the Iberian Peninsula. Although the genus Crupina comprises two species in Spain, only one is known in the Western Hemisphere. This study compiled distribution maps for both C. vulgaris and C. crupinastrum in the Iberian Peninsula using 939 collections from 17 herbaria in Spain and 154 locations recorded in the literature (floras, botanical explorations, and phytosociological studies). Crupina vulgaris is more com- mon, with a distribution occupying most of the eastern half of the Peninsula, reaching north to the southern slopes of the Pyrenees Mountains which form the boundary between Spain and France. It is rare in the western third of the Peninsula, including Portugal, and is absent on the Balearic Islands. Found throughout the southeastern quadrant of the Peninsula and Mallorca in the Balearic Islands, Crupina crupinastrum is most abundant in the Andalusian provinces. In southeastern Spain where their distributions overlap, C. crupinastrum is more prevalent than C. vulgaris. Detailed distribution maps in the native range of invasive species are useful for understanding biological invasions, comparisons of native and introduced habitats, and searches for potential biological control agents. RESUMEN Crupina vulgaris es una planta de origen mediterraneo, de entre las 1045 especies naturalizadas en California, que se comporta como invasora y cuenta con poblaciones, relativamente recientes, en los condados de Sonoma y Modoc. Los estudios moleculares han demostrado que las cinco poblaciones norteamericanas derivan de 3 0 mas introducciones procedentes de la Peninsula Ibérica. Aunque el género Crupina comprende dos especies en Espana, solamente una es conocida en el hermisferio occidental. En este estudio se presentan los mapas de distribuci6n de C. vulgaris y C. crupinastrum en la Peninsula Ibérica, obtenidas tras la revision de 939 recolecciones encontradas en 17 herbarios de Espana y 154 referencias bibliograficas (floras, catalogos floristicos, y estudios fitosociolé6gicos). Crupina vulgaris es la especie mas comun, con una distribuci6n que ocupa la mitad oriental de la Peninsula, alcanzando hacia el Norte las vertientes meridionales de los Pirineos. Es rara en el tercio oeste, incluyendo Portugal, y esta ausente en las islas Baleares. Presente en todo el cuadrante suroriental de la Peninsula y en Mallorca (islas Baleares), Crupina crupinastrum es mas abundante en las provincias andaluzas. En el Sureste de Espana, donde ambas distribuciones se solapan, C. crupinastrum prevalece sobre C. vulgaris. Los mapas de distribuci6n detallados de las especies invasoras en su zona de origen, son utiles para comprender las invasiones, la comparaci6n entre habitat autdctonos y aléctonos, y las investigaciones para buscar poten- ciales agentes de control bioldégico. Key words: Crupina crupinastrum, Crupina vulgaris, invasive species, weed distribution Among the 1045 alien plant species naturalized (17.5% of the flora) in California (Randall et al. 1998), Crupina vulgaris Cass. (Asteraceae: Cyna- reae) is a relatively recent addition. Rejmanek and Randall (1994) indicated that it was very likely in- troduced to California within the past 25 years. This contrasts sharply with numerous other Mediterra- nean species introduced in the Spanish and Mexi- can periods (1769-1848) (Bossard et al. 2000). When Crupina vulgaris was discovered in Idaho in 1968, it was reported in Madrofio as a species new * Corresponding author. to North America (Stickney 1972). Under the Fed- eral Noxious Weed Act of 1974 (Public Law 93 629), Crupina vulgaris was included on the Federal Noxious Weed List as a new invader. Both Crupina vulgaris and its congener Crupina crupinastrum (Moris) Vis. were among 1200 species designated as economically important foreign weeds posing potential problems in the United States (Reed 1977). A search of the interception records for Fed- eral Noxious Weeds by USDA APHIS PPQ from 1983 through 1998 showed that Crupina vulgaris has not been intercepted crossing the U.S. borders since it was listed and records kept on weed inter- 138 ventions (Polly Lehtonen, USDA APHIS, personal communication). Although the first flora in California to include Crupina vulgaris was The Jepson Manual (Hick- man 1993), it was first discovered in California in 1976 at Santa Rosa, Sonoma County (unpublished CDFA reports, Miller and Thill 1983). It was sub- sequently discovered in 1984 at Lake Chelan, Che- lan County, Washington (Alverson and Arnett 1986), and in 1987 at Dry Creek, Umatilla County, Oregon (Couderc-LeVaillant and Roché 1993). The Sonoma County population was declared eradicated by 1982 (Miller and Thill 1983), but it was redis- covered in 1989 about 1 km distant from the orig- inal infestation (unpublished CDFA records, Davis and Sherman 1991). In 1990 another new popula- tion was reported near Adin, Modoc County, Cali- fornia (unpublished CDFA records, Couderc-Le- Vaillant and Roché 1993). Thus, five infestations in four western States were detected between 1968 and 1990. Crupina vulgaris is inconspicuous due to its small, delicate stature and extremely difficult to detect when populations are sparse. All six original discoveries of C. vulgaris in the western United States were made by professional botanists or weed scientists; none were reported by casual observers or landowners. Multiple introductions have been suspected for the invasion (Couderc-Le Vaillant and Roché 1993; Patterson and Mortensen 1985), although a large propagule hinders long distance dispersal (Roché and Thill 2001). Since the previous publications in Madrono, earlier collections of C. vulgaris in North America have been reported from Massachusetts (Sorrie and Somers 1999). Specimens at the Har- vard University Herbaria collected by C.E. Perkins in 1877 and 1879 from Boston and South Boston Flats, Suffolk County, Massachusetts (NEBC, W. Kittridge personal communication), indicate that C. vulgaris was among the numerous species intro- duced in ship’s ballast from seaports in the Medi- terranean region. Based on its absence in current floras, C. vulgaris failed to establish in the north- eastern United States, and apparently arrived inde- pendently in western North America. Recent mo- lecular studies (RAPD) revealed that the five cur- rent populations derived from three or more intro- ductions from the Iberian Peninsula (Garnatje et al. 2002). Although the genus Crupina is represented by two species in the Iberian Peninsula and Bale- aric Islands, C. crupinastrum and C. vulgaris, only one has been reported in North America. The objective of the study was to compile an accurate distribution map for Crupina in the part of its native range where its North American popula- tion founders originated. Such a map would serve as a foundation for further investigations, such as ecological studies comparing native and introduced habitats, including behavior of the invader, and fac- tors that contribute to differences in species re- sponse in the two hemispheres. A distribution map MADRONO [Vol. 49 is also useful for searches for potential biological control agents. Floras provide general distribution information that is inadequate for these purposes. For example, Flora Europaea indicates that C. vul- garis grows on “dry grassland and stony slopes”’ in 17 countries (Amaral Franco 1976), which leads one to believe that it can be easily encountered any- where in these habitats in southern Europe, as far north as west central France and the southern Ukraine. In fact, much of this distribution is based on centuries-old collection records, some of which represent localized populations that failed to persist under changing land use patterns in the past 100 years. In our study we included the more ruderal (within the Mediterranean region) congener, C. cru- pinastrum, in order to provide supporting infor- mation for inferences about plant migration and the invasion process. METHODS A complete listing of all recorded locations for both species of Crupina in the Iberian Peninsula was compiled from two types of sources: 1) liter- ature citations of locations from floras, botanical explorations, and phytosociological studies and 2) herbarium specimens from 17 herbaria in Spain (listed in the acknowledgments, with institutional abbreviations from Holmgren et al. 1990). Because the two species closely resemble each other, each herbarium sheet was examined and annotated. On mature specimens, verification was based on cyp- sela characters, while immature specimens and oth- ers without fruits were verified using trichome char- acters (Couderc-LeVaillant 1984). Maps were pre- pared using software (CYANUS) based on 10 km UTM grid square. RESULTS The distributions of C. vulgaris and C. crupinas- trum are shown on maps in Figs. 1 and 2, respec- tively. Of the 939 herbarium specimens examined, 572 were assigned to C. vulgaris and 367 to C. crupinastrum. The maps include 134 citations from the literature for which there was no doubt about the identification. An additional 120 literature ci- tations for C. vulgaris were not included on the maps because they lacked corroborating herbarium vouchers and were from regions where the distri- bution of C. crupinastrum overlapped with that of C. vulgaris. In some provinces there were locality descriptions for which UTM coordinates could not be determined. A complete record of herbarium la- bel and literature citations will be published in Spain (Gamarra and Roché 2002). Crupina vulgaris In the Iberian Peninsula, C. vulgaris is the more common species, with a distribution occupying most of the eastern half of the Peninsula (Fig. 1). All of the phytosociology literature and plant dis- 2002] ‘Dizerress = ee GAMARRA AND ROCHE: CRUPINA IN THE IBERIAN PENINSULA 139 e EDS Ne 1 e s ses 4 al ‘ oN codes Sa Na 36° Fic. 1. Distribution of Crupina vulgaris in the Iberian Peninsula, each symbol indicates documented presence within a 10 km UTM grid square. Dotted lines are provincial boundaries; refer to Fig. 2 for names. tribution records indicate that C. vulgaris is not found in forested areas and in subalpine grasslands, as well as areas of siliceous soils, which tend to be coarse and acidic. It does not occur on the Balearic Islands. The most northerly populations of C. vul- garis reach the southern slopes of the Pyrenees and the Pre-Pyrenean Mountains. In the western third of the peninsula its presence is sparser and almost always tied to calcareous substrates, which are in- frequent in this region. In Portugal it is very rare, limited to certain populations near the locality of Elvas, very close to the border with Spain, and far- ther away, in Sezimbra, but always on calcareous soils. Using locations from herbarium labels dating from the 1970’s, one of the authors (Roché) and a colleague searched all suitable habitat in the Elvas locality in 1999 and failed to find a single plant. If C. vulgaris is still present there, it is extremely scarce. According to Rivas Martinez et al. (1990), this species appears in all the chorologic provinces, al- though in the north and northeastern provinces of Cantabro-Atlantica and Gaditano-Onubo-Algar- viense, it occurs only in isolated locations. As stat- ed earlier, in the history of Spanish botanical sci- ence, it has never been found in the Balearic prov- ince. The elevation within its distribution ranges be- tween 100 and 1200 (rarely 1500) m, reaching the major territory of the Sierra de Segura and the Pyr- enees, but it is never found in the high mountains. It prefers basic soils, principally calcareous soils derived from limestone or clay soils rich in bases originating from evaporites (formed by the evapo- ration of brackish water), including substrates rich in gypsum; and is only rarely found over schist or slate. It is reported from rangeland, dry grazed ar- eas, low matorral (e.g., thyme fields), open ever- green oak woodlands, and clearings in deciduous oak forests. It is also found along roadsides and on the margins of perennial crops such as vineyards or olive groves where they border appropriate native habitat. Crupina crupinastrum This species primarily inhabits the southeastern quadrant of the peninsula and the Balearic Islands, 140 MADRONO [Vol. 49 Pizerress Fic. 2. 42° | | | Distribution of Crupina crupinastrum in the Iberian Peninsula, each symbol indicates documented presence within a 10 km UTM grid square. Abbreviations for names of provinces: Alicante (A), Albacete (Ab), Almeria (Al), Asturias (O), Avila (Av), Badajoz (Ba), Barcelona (B), Burgos (Bu), Caceres (Cc), Cadiz (Ca), Cantabria (S), Castellon (Cs), Cordoba (Co), Ciudad Real (CR), Cuenca (Cu), Gerona (Ge), Granada (Gr), Guadalajara (Gu), Huelva (H), Huesca (Hu), Jaen (J), La Corufia (C), La Rioja (Lo), Ledn (Le), Lerida (L), Lugo (Lu), Madrid (M), Malaga (Ma), Murcia (Mu), Navarre (Na), Orense (Or), Palencia (P), Pontevedra (Po), Salamanca (Sa), Sevilla (Se), Segovia (Sg), Soria (So), Tarragona (T), Teruel (Te), Toledo (To), Valencia (V), Valladolid (Va), Vizcaya (Vi), Guipuzcoa (SS), Zamora (Za), Zaragoza (Z). and is most abundant in the Andalusian provinces (Almeria, Cadiz, Cordoba, Granada, Huelva, Jaén, Malaga, Sevilla) (Fig. 2). In contrast to the northern presence of C. vulgaris, C. crupinastrum only ex- tends as far north as the provinces of Segovia and Burgos, where it is rarely encountered and could be interpreted as isolated individuals occurring as ephemeral introductions. In the Balearic Islands, it is found only on Mallorca. It does not appear on the Pitiusas Islands (the southwest islands within the Balearic group), which would be an extension of the levantan populations. Toward western Iberia, it is very rare and we found only isolated occur- rences in the provinces of Sevilla (Castilleja de Guzman) and Caceres (Guadalupe), which did not extend as far as Portugal. According to Rivas Martinez et al. (1990), this species appears principally in the provinces Bética and Castellano-Maestrazgo-Manchega, with some presence in the provinces Murciano-Almeriense and Balear, and is very rare in the Luso-Extrema- durense (Portugal, Caceres and Badajoz). The presence of Crupina crupinastrum is linked to basic soils, principally substrates of limestone and evaporites, although some populations in the province of Cérdoba grow over schists. The ele- vation ranges between 100 and 1500 m, very rarely exceeding this limit, but occasionally doing so in the more southern mountains as in Gador and in the Sierra Nevada. It appears on the sides of roads, grazed lands, pastures, low matorral, and open ev- ergreen oak woodlands. DISCUSSION Both species of Crupina share the southeastern quadrant of the Iberian Peninsula, in some cases growing in mixed communities. In this region C. 2002] crupinastrum is much more abundant than C. vul- garis. However, nowhere in the Iberian Peninsula would one describe either species as abundant in the plant communities where they occur. It is note- worthy that the more ruderal C. crupinastrum, which appears more frequently in disturbed sites (e.g., roadsides) than C. vulgaris, was not the spe- cies introduced in North America. Because cypselas of the two species are the same size with the same pappus characteristics, differing only in shape at the point of attachment (see illustrations in Reed 1977), this anomaly suggests that the invasion founders originated from locations where C. crupinastrum does not grow along with C. vulgaris. On sites supporting Crupina in the Iberian Pen- insula, grazing of sheep and goats is the primary land use, especially historically. In the Mediterra- nean region, Crupina has been identified among low chaparral species which maintain populations by epizooic transport associated with herds of do- mestic sheep and goats (Schmida and Ellner 1983). Transhumanant herds of sheep were likely the mi- gratory vectors responsible for dispersing and maintaining isolated ephemeral populations in the Iberian Peninsula. Well established trails (cafiadas) connect summer and winter pastures (Montserrat and Fillat 1990), which may be as close as moun- tain grasslands with adjacent valleys, or extend nearly the entire north-south distance of Spain (Mangas 1992). The distribution of C. vulgaris far to the north of C. crupinastrum in the Iberian Peninsula is es- pecially significant because the Pyrenean and Pre- Pyrenean Mountain region was the source of major immigration to the United States of laborers for the sheep industry through the early 1970’s. Basque sheepherders were legendary in the western U.S. For example, in 1970 about 90% of the 1700 men under contract to the Western Range Association were Basques (Lane and Douglass 1985). Although the Spanish government required equal immigra- tion opportunities for all Spanish nationals, it was primarily due to improving economic conditions in the Basque Country in the 1960’s and 1970’s that the numbers of Asturians, Leonese, Castillians and Andalusians swelled the herder ranks (Lane and Douglass 1985). After that time, this link with rural Spain ended when the range association shifted its herder recruitment efforts to Latin America and Mongolia. Despite the recent (1968—1990) discoveries of C. vulgaris in western North America, it is probable that it arrived decades earlier. Arriving in small numbers to remote areas, with slow growing colo- nies of inconspicuous individuals, it is not surpris- ing that it could escape detection for long periods of time. By showing where the distributions of C. vulgaris and C. crupinastrum overlap and diverge, and where the distribution of C. vulgaris overlaps a region that was home to numerous immigrants to rangelands of the western United States, these dis- GAMARRA AND ROCHE: CRUPINA IN THE IBERIAN PENINSULA 141 tribution maps provide a resource for elucidating the case history of how C. vulgaris became an in- vader in the Western Hemisphere, a chronicle that has not yet been fully revealed. They also serve as a reference for future studies concerning ecology and potential control. ACKNOWLEDGMENTS The following herbaria in Spain graciously loaned or allowed us to review their materials: Sociedad de Ciencias Aranzadi, San Sebastian (ARAN): Institut Botanic de Bar- celona, Barcelona (BC); Universitat de Barcelona (Depar- tament de Biologia Vegetal), Barcelona (BCC); Univer- sitat de Barcelona (Facultat de Farmacia), Barcelona (BCF); Universidad de Cérdoba, Cérdoba (COFC); Uni- versidad de Granada, Granada (GDA): Universidad de Granada, Granada (GDAC); Universitat de Girona, Girona (HGI); Instituto Pirenaico de Ecologia, Jaca (JACA): Co- legio Universitario Santo Reino, Jaén (JAEN); Real Jardin Botanico, Madrid (MA); Universidad Complutense (Cien- cias Biolégicas), Madrid (MACB); Universidad Complu- tense (Farmacia), Madrid (MAF); Universidad de Malaga, Malaga (MGC); Universidad de Sevilla, Sevilla (SEV): Universitat de Valéncia, Valencia (VAL); and Museo de Ciencias Naturales de Alava, Vitoria (VIT). We thank the following individuals for their generous sharing of their time and knowledge: Daniel Gémez, Federico Fillat, Mi- kel Lorda, Pedro Uribe, Mauricio Velayos, José Pizarro, Josep Vicens and Antonio Sanchez-Cuxart. The manu- script was reviewed by K. L. Chambers. C. Roché extends special thanks to Josep Montserrat, for his encouragement and his suggestions of contacts for the project. LITERATURE CITED ALVERSON, E. AND J. ARNETT. 1986. Plant life of the North Cascades: Lake Chelan-Sawtooth Ridge, Stehekin Valley and Glacier Peak. Douglasia Occasional Paper, Vol. 2. Washington Native Plant Society, Seattle, WA. AMARAL FRANCO, J. 1976. Crupina. P. 301 in T. G. Tutin et al. (eds.), Flora Europaea, Vol. 4. Cambridge Univ. Press, London, U.K. BOSSARD, C. C., J. M. RANDALL, AND M. C. HOSHOVSKY (eds.). 2000. Invasive plants of California’s wildlands. University of California Press, Berkeley, CA. COUDERC-LEVAILLANT, M. 1984. L-amphiploidie dans le genre Crupina DC. Essai de systématique synthéti- que. Thése Docteur Es-Sciences Naturelles, Univ. Paris-Sud, Orsay, France. AND C. T. ROCHE. 1993. Evidence of multiple in- troduction of Crupina vulgaris in infestations in the western United States. Madrono 40:63-—65. Davis, L. H. AND R. J. SHERMAN. 1991. Crupina vulgaris Cass. (Asteraceae: Cynareae), established in Sonoma County, California, at Annadel State Park. Madrono 38:296. GAMARRA, R. AND C. T. ROCHE. 2002. Cartografia Coro- l6gica Ibérica. Aportaciones. Botanica Complutensis. Universidad Complutense, Madrid, Spain. GARNATIE, T., R. VILATERSANA, C. T. ROCHE, N. GARCIA- JACAS, A. SUSANNA, AND D. C. THILL. 2002. Multiple introductions from the Iberian Peninsula responsible for invasion of Crupina vulgaris Cass. in western North America. New Phytologist 154:419—428. HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. 142 MADRONO HOLMGREN, P. K., N. H. HOLMGREN, AND L. C. BARNETT. 1990. Index herbariorum, 8th ed. New York Botanical Garden, Bronx, NY. LANE, R. H. AND W. A. DouGLass. 1985. Basque sheep- herders of the American West. University of Nevada Press, Reno, NV. MANGAS, J. M. 1992. Vias pecuarias. Cuadernos de la tras- humancia, No. 0, Instituto Nacional para la Conser- vacion de la Naturaleza, Madrid, Spain. MILLER T. AND D. THILL. 1983. Today’s weed: common crupina. Weeds Today 14:10—11. MONTSERRAT, P. AND FE FILLAT. 1990. The systems of grassland management in Spain. Pp. 37-70 in A. Breymeyer (ed.), Managed grasslands, ecosystems of the world. Elsevier, Amsterdam, Netherlands. PATTERSON, D. E. AND D. A. MorTENSEN. 1985. Effects of temperature and photoperiod on common crupina (Crupina vulgaris). Weed Science 33:333-339. RANDALL, J. M., M. REJMANEK, AND J. C. HUNTER. 1998. Characteristics of the exotic flora of California. Fre- montia 26(4):3—-12. REED, C. E 1977. Economically important foreign weeds. Ag. Handbook No. 498. U.S. Govt. Printing Office, Washington, DC. [Vol. 49 REJMANEK, M. AND J. M. RANDALL. 1994. Invasive alien plants in California: 1993 summary and comparison with other areas in North America. Madrono 41:161— Wa Tle RivAs MARTINEZ, S., P. CANTO, E FERNANDEZ GONZALEZ, C. NAVARRO, J. M. PIZARRO, AND D. SANCHEZ MATA. 1990. Biogeografia de la Peninsula Ibérica, Islas Ba- leares y Canarias. X Jornadas de Fitosociologia, Gra- nada, Spain. ROCHE, C. T. AND D. C. THILL. 2001. Biology of common crupina and yellow starthistle, two Mediterranean winter annual invaders in western North America. Weed Science 49:439—447. SCHMIDA, A. AND S. ELLNER. 1983. Seed dispersal on pas- toral grazers in open Mediterranean chaparral, Israel. Israel Journal of Botany 32:147—159. SORRIE, B. A. AND P. Somers. 1999. The vascular plants of Massachusetts: a county checklist. Massachusetts Division of Fisheries and Wildlife, Natural Heritage and Endangered Species Program, Westborough, MA. STICKNEY, P. E 1972. Crupina vulgaris (Compositae: Cynareae), new to Idaho and North America. Madro- no 21:402. MaApbroNo, Vol. 49, No. 3, pp. 143-144, 2002 HYMENOCLEAS ARE AMBROSIAS (COMPOSITAE) JOHN L. STROTHER University Herbarium, University of California, Berkeley, CA 94720-2465 strother @uclink4.berkeley.edu BRUCE G. BALDWIN Jepson Herbarium and Department of Integrative Biology, University of California, Berkeley, CA 94720-2465 ABSTRACT Inclusion of Hymenoclea within the taxonomic circumscription of Ambrosia necessitates new combi- nations: Ambrosia monogyra, A. xplatyspina, A. salsola, A. salsola var. fasciculata, and A. salsola var. pentalepis. Key words: Ambrosia, Compositae, Hymenoclea After review of similarities and differences be- tween and among species of Hymenoclea Torrey & A. Gray ex A. Gray and Ambrosia Linnaeus, es- pecially with regard to restriction sites in chloro- plast DNAs, Miao et al. (1995) concluded that the two species of Hymenoclea do not constitute a clade, are separately allied to franserioid members of Ambrosia, and are better included in Ambrosia than maintained as a distinct genus. They listed Hy- menoclea as a synonym of Ambrosia Linnaeus subg. Franseria (Cav.) Miao et al., Pl. Syst. Evol. 194:252, 1995. Baldwin et al. (1996) documented natural hy- bridization between Hymenoclea salsola Torrey & A. Gray ex A. Gray and the franserioid species Am- brosia dumosa (A. Gray) W. W. Payne and between H. salsola and A. ambrosioides (Cavanilles) W. W. Payne, another franserioid species. They noted “.. . normal pairing of chromosomes in interspecific hy- brids”’ (1.e., between species of Ambrosia and Hy- menoclea) as indicating ““Close genetic similarity ” of parental species and went on to say, ““Hy- bridization between species of Ambrosia and Hy- menoclea may reflect inadequacy of the long-stand- ing generic classification of Ambrosiinae ... .” We have considered the findings of Miao et al. (1995) and Baldwin et al. (1996) and we are con- vinced that hymenocleas should be treated within the taxonomic circumscription of Ambrosia: Ambrosia monogyra (Torrey & A. Gray ex A. Gray) Strother & B. G. Baldwin, comb. nov. Basionym: Hymenoclea monogyra Torrey & A. Gray ex A. Gray, Mem. Amer. Acad. Arts ns. 4: 79. 1849. Syntypes: “‘Along the valley of the Gila, Lieut. Emory. Also at ‘Ojito,) New Mexi- co? Dr. Gregg.’ Peterson and Payne (1973, p. 253-254) cited the Emory collection at NY as type and thereby effected lectotypification. Ambrosia salsola (Torrey & A. Gray ex A. Gray) Strother & B. G. Baldwin, comb. nov. Basionym: Hymenoclea salsola Torrey & A. Gray ex A. Gray, Mem. Amer. Acad. Arts n.s. 4:79. 1849. Type: California, ““Sandy, saline uplands near the Mojave River, ..., Fremont.’’ Lectotype (Peter- son and Payne 1973, p. 254): 1844, Frémont 400 (NY). Ambrosia salsola (Torrey & A. Gray ex A. Gray) Strother & B. G. Baldwin var. fasciculata (A. Nelson) Strother & B. G. Baldwin, comb. nov. Basionym: Hymenoclea fasciculata A. Nelson var. fasciculata. [cf. Hymenoclea fasciculata A. Nelson, Bot. Gaz. 37:270. 1904. Type: Nevada, ““Kernan,”’ 29 Apr 1902, L. N. Goodding 662 (NY). The variety fasciculata dates from publi- cation of Hymenoclea fasciculata A. Nelson var. patula A. Nelson, Bot. Gaz. 47:431. 1909. Type: Nevada, “‘“Moapa,”’ 8 Apr 1905, L. N. Goodding 2178 (RM). Peterson and Payne (1973, 1974) treated the two types, Goodding 662 and 2178, as convarietal. Initially, Peterson and Payne named that variety Hymenoclea salsola Torrey & A. Gray ex A. Gray var. fasciculata (A. Nelson) K. M. Peterson & W. W. Payne (Brittonia 25:255. 1973). Under the applicable Code (Stafleu et al. 1972) in 1973, Peterson and Payne should have used the varietal epithet “‘patula’’ because auto- nyms were “not to be taken into consideration for purposes of priority.”’ Peterson and Payne subsequently renamed that same variety Hymen- oclea salsola Torrey & A. Gray ex A. Gray var. patula (A. Nelson) K. M. Peterson & W. W. Payne (Brittonia 26:397. 1974), which is now an illegitimate name because under the current Code (Greuter et al. 2000), the autonymic varietal name has priority at varietal rank. ] Ambrosia salsola (Torrey & A. Gray ex A. Gray) Strother & B. G. Baldwin var. pentalepis (Ryd- berg) Strother & B. G. Baldwin, comb. nov. Bas- ionym: Hymenoclea pentalepis Rydberg in N. L. 144 MADRONO Britton et al., N. Amer. Fl. 33:14. 1922. Type: Arizona, “‘Pima Canon,”’ 10 Apr 1901, D. Grif- fiths 2630 (NY) = Hymenoclea salsola Torrey & A. Gray ex A. Gray var. pentalepis (Rydberg) L. D. Benson, Amer. J. Bot. 30:631. 1943. Hymenoclea hemidioica A. Nelson, Amer. J. Bot. 25:117. 1938. Syntypes: Arizona, Mohawk Mountains, 29 Mar 1935, A. Nelson 1340 and 1341 (RM?, not seen). We use the name Ambrosia xplatyspina (Sea- man) Strother & B. G. Baldwin, comb. nov. [Bas- ionym: Hymenoclea xplatyspina Seaman, Madrono 23:111. 1975, pro sp.], for hybrids between A. du- mosa and A. salsola. We believe the name Ambrosia sandersonii S. L. Welsh, Rhodora 95:396. 1993[1994] [=Hymeno- clea sandersonii (S. L. Welsh) N. H. Holmgren in A. Cronquist et al., Intermount. Fl. 5:473. 1994], also refers to hybrids, perhaps to hybrids between A. eriocentra (A. Gray) W. W. Payne and A. salsola (Baldwin et al. 1996). We treat Welsh’s name as Ambrosia xsandersonii S. L. Welsh, pro sp. [Vol. 49 ACKNOWLEDGMENTS We thank D. J. Keil, T. K. Lowrey, R. L. Moe, and A. R. Smith for helpful comments on early drafts of this pa- per. LITERATURE CITED BALDwIn, B. G., D. W. KyuHos, S. N. MARTENS, FE C. Va- SEK, AND B. L. WEsSA. 1996. Natural hybridization between species of Ambrosia and Hymenoclea sal- sola (Compositae). Madrofo 43:15—27. GREUTER, W., ET AL. 2000. International code of botanical nomenclature. Regnum Vegetabile 138:v—xvili, 1— 474. Miao, B., B. TURNER, B. SIMPSON, AND T. MABRY. 1995. Chloroplast DNA study of the genera Ambrosia s.1. and Hymenoclea (Asteraceae): systematic implica- tions. Plant Systematics and Evolution 194:141—255. PETERSON, K. M. AND W. W. PAYNE. 1973. The genus Hy- menoclea (Compositae: Ambrosieae). Brittonia 25: 243-256. AND . 1974. Erratum: the correct name for the appressed-winged variety of Hymenoclea salsola (Compositae: Ambrosieae). Brittonia 26:397. STAFLEU, FE A., ET AL. 1972. International code of botan- ical nomenclature. Regnum Vegetabile 82:7—426. Mapbrono, Vol. 49, No. 3, pp. 145-149, 2002 THE FLORA OF TORO ISLET AND NOTES ON GUADALUPE ISLAND, BAJA CALIFORNIA, MEXICO JON P. REBMAN San Diego Natural History Museum, P.O. Box 121390, San Diego, CA 92112-1390, USA jrebman @sdnhm.org THOMAS A. OBERBAUER County of San Diego, Department of Planning and Land Use, 5201 Ruffin Rd. Suite B5, San Diego, CA 92123, USA Jose Luts LEON DE LA Luz Centro de Investigaciones Bioldégicas del Noroeste, Apdo. Postal 128, La Paz, 23000, Baja California Sur, México ABSTRACT During a natural history expedition to Guadalupe Island and its adjacent islets in June of 2000, the previously unexplored islet, Toro, was botanically surveyed. The flora of this islet was found to have 32 species and one putative interspecific hybrid. This diversity represents 30 dicots and 2 monocots, in 22 plant families. Eighteen of the plant taxa are endemic to the Guadalupe Island group, resulting in a 56.3% endemism in the islet’s flora. A few native plant communities were observed on the islet, within which only three plant taxa are obviously exotic, rare in occurrence, and presumably introduced by seabirds. The botanical data obtained from this undisturbed islet helps us to fill in the missing pieces about the overall flora of Guadalupe Island and its adjacent islets especially in relation to the ecological devastation caused by feral goats on the main island. Brief notes on other floristic components of Guadalupe Island and its islets, and new plant records collected during the expedition are also presented. RESUMEN Durante una expedicion para estudiar la historia natural de la Isla Guadalupe e islotes adyacentes, en junio de 2000, inventariamos la flora del islote Toro, que nunca habia sido explorado. En este islote encontramos 32 especies de plantas y un aparente hibrido interespecifico, incluyendo 30 dicotiledonas y 2 monocotiledonas, de 22 familias. Dieciocho de los taxa son endémicos a la Isla Guadalupe e islotes adyacentes, por lo tanto Toro tiene una tasa de endemismo de 56.3%. Se encontraron algunas comunidades de plantas nativas, en las que solamente 3 taxa eran ex6ticas, poco abundantes, probablemente introducidas por aves marinas. La informacion botanica de este islote sin disturbio puede ayudarnos a llenar las piezas faltantes de la flora de Guadalupe, especialmente en relaci6n con la devastaci6n ecolégica causada por cabras en la isla principal. Ademdas, en este articulo presentamos notas breves sobre otros componentes floristicos de la Isla Guadalupe y sus islotes, y nuevos registros de plantas colectadas durante la expe- dicion. Key words: Guadalupe Island, Baja California, Mexican flora, expedition, endemism The Guadalupe Island Expedition in June of 2000 involved 16 scientists from the U.S.A. and Mexico representing arachnology, botany, conser- vation biology, entomology, marine ecology, orni- thology, and phycology. This expedition was or- ganized by the San Diego Natural History Museum and funded by the Biotic Surveys and Inventories section of the National Science Foundation (Grant No. 0074462). The primary emphases of this bi- national, multidisciplinary endeavor were to ob- serve, record, and collect natural history informa- tion on the biodiversity of Guadalupe Island and the previously unexplored Toro Islet. The botanists of the expedition were Dr. José Luis Leon de la Luz of the Centro de Investigaciones Bioldgicas del Noroeste, Tom Oberbauer of the Department of Planning and Land Use for the County of San Di- ego, and Dr. Exequiel Ezcurra, and the senior au- thor from the San Diego Natural History Museum. The biological data obtained on this trip provides us with additional biodiversity information on all of the terrestrial areas of Guadalupe Island and its islets and will be used for making conservation de- cisions for the island and evaluating the impacts of introduced species such as feral goats and cats. The floristic information from the expedition can be used to supplement our current knowledge on the entire Guadalupe Island flora as presented by Mo- ran (1996). THE ISLAND ENVIRONMENT Guadalupe is an oceanic island located approxi- mately 260 km off the Pacific coast of the Baja 146 California peninsula, in northwestern Mexico. The island’s general geographic position is 29°03'N lat- itude and 118°17’W longitude. Guadalupe Island is about 36 km long on its N-S axis and 12 km wide on the E-W axis, with an approximate surface area of 250 km/?. The island is the peak of a seamount, which may have originated from several eruptive episodes, with the oldest exposed rocks being dated around 7 million years old (Moran 1996). There are three islets off of the southern end of the main is- land: Negro (30 m in elevation), Toro (220 m), and Zapato (190 m). Each islet has approximately 1 km? of surface area. The climate of the island is maritime and heavily influenced by the cold California Current, with its characteristic features of wind, fog, and winter rain- fall. A meteorological station on the southern end of Guadalupe (the driest area of the island) indi- cates almost 120 mm of rainfall annually and a mean monthly temperature of 17—19°C that is rel- atively stable throughout the year. It should be not- ed that the annual precipitation value given above does not include the contribution of moisture from fog condensation, which is a common event on Guadalupe Island. Floristically, the island is considered an “‘outli- er’ of the California Floristic Province (Moran 1996) since it is composed of many native plant species either disjunct from this vegetation type on the mainland or on other islands, or endemic spe- cies with northern affinities. Although the island is home to Mexico’s only populations of many plant species from the California Floristic Province, its most striking attribute is its endemism, which oc- curs in many biological groups. According to Mo- ran (1996) in respect to plants, almost 22% of the native species are endemic, including two mono- specific genera. Although there have been 220 dif- ferent plant taxa documented over time on Guada- lupe Island and its islets, the activities of feral goats released in the early 19th century by whalers have devastated most of the main island’s flora. This im- pact on the flora is not only from the direct browse of goats, but also from the damage to the insular substrates as a result of soil erosion. At present, the main island’s original flora and natural plant com- munities have practically vanished. It is estimated (Moran 1996) that at least 26 native plant species could now be extinct, including one endemic, monospecific genus and many other plant species seem to be on the threshold. To add to the problem, the vegetation is now dominated mainly by weedy species; 62 exotic plant taxa have been recorded on the island, of which many are quite aggressive and capable of competing for niches and displacing na- tive species. Toro (also known as Islote de Adentro or Inner Islet) is a small islet that lies directly off of the southern end of Guadalupe Island, situated between the main island and another southern islet, Zapato. Toro Islet is a dome-shaped rock that reaches ap- MADRONO [Vol. 49 proximately 500—700 feet (165-220 m) in eleva- tion. The islet has very steep, vertical walls that arise out of the water at an angle of almost 90 de- grees and that have, heretofore, prohibited its ex- ploration. However, during this expedition the sci- entists used a helicopter to facilitate landing and exploration on the islet. On its top, Toro has a small basin in its center with a high western ridge. A narrow ridge extends to the north and drops into a steep slope facing the main island. There are two types of rocky substrates that exist on this volcanic islet. One type of substrate is a rocky, fractured basaltic material with darker coloration and the oth- er is a tan, hard solid rock with characteristics of andesite. Plant species occur on both substrates but most seem to prefer the broken basaltic materials. THE FLORA OF TORO The known flora (see Table 1) of Toro Islet that was documented during the expedition consists of 32 species and | putative hybrid. This diversity rep- resents 30 dicots and 2 monocots, in 22 plant fam- ilies. Eighteen of the taxa are endemic to Guada- lupe Island, resulting in a 56.3% rate of endemism in the islet’s flora. The Asteraceae are the best rep- resented on the islet with 5 genera, 6 species and one putative interspecific hybrid in the genus He- mizonia. Other families such as Malvaceae, Scrop- hulariaceae, and Fabaceae are represented with two genera. Only Cryptantha, Hemizonia, and Mesem- bryanthemum are represented with two species in the same genus. Only three plant taxa (Hordeum murinum ssp. glaucum, Mesembryanthemum crystallinum, and M. nodiflorum) are obviously exotic. As of yet, their populations on the islet are small and their occur- rences quite rare. It is likely that they are rather recent introductions, which presumably arrived by seabirds. In fact, Hordeum seems to be currently confined to the steep northern slope of the islet in an area near Western Gull (Larus occidentalis) nests. Of the 32 plant species found on the islet, eleven are succulent in nature with fleshy leaves, stems, or both, and are represented by species in eight dif- ferent plant families (Rebman 2001). Six of these succulent taxa are endemic to the Guadalupe Island and its adjacent islets. One of the leaf succulent species, Baeriopsis guadalupensis belongs to an en- demic, monotypic genus in the Sunflower family (Asteraceae). The vegetation of Toro Islet is best described as a maritime, succulent scrub and is similar to that found on the other southern islets, Zapato and Ne- gro. This type of vegetation probably dominated most of the southern portions of the main island as well, but is mostly extirpated now by the impacts of feral goats. This vegetation is dominated by Azri- plex barclayana, Cistanthe guadalupensis, Euphor- bia misera, Spergularia macrotheca var. talinum, 2002] TABLE 1. of plant specimens made by Rebman et al. Plant taxon (family) Atriplex barclayana (Benth.) D. Dietr. (Chenopodiaceae) Baeriopsis guadalupensis J. T. Howell (Asteraceae) Castilleja fruticosa Moran (Scrophulariaceae) Cistanthe guadalupensis (Dudley) Carolin in Hershkovitz (Portulacaceae) Coreopsis gigantea (Kellogg) H. M. Hall (Asteraceae) Cryptantha foliosa (Greene) Greene (Boraginaceae) Cryptantha maritima (Greene) Greene var. maritima (Boraginaceae) Dichelostemma capitatum Alph. Wood ssp. capitatum (Themidaceae) Dudleya guadalupensis Moran (Crassulaceae) Erysimum moranii Rollins (Brassicaceae) Eschscholzia palmeri Rose (Papaveraceae) Euphorbia misera Benth. (Euphorbiaceae) Galvezia speciosa (Nutt.) A. Gray (Scrophulariaceae) Hemizonia greeneana Rose ssp. greeneana (Asteraceae) Hemizonia hybrid (Asteraceae) Hemizonia palmeri Rose (Asteraceae) Hordeum murinum L. ssp. glaucum (Steud.) Tzvelev. (Poaceae) Lavatera lindsayi Moran (Malvaceae) Lomatium insulare (Eastw.) Munz (Apiaceae) Lotus argophyllus (A. Gray) Greene ssp. ornithopus (Greene) Raven (Fabaceae) Lupinus niveus S. Watson (Fabaceae) Lycium californicum Nutt. (Solanaceae) Mammillaria blossfeldiana Boed. var. shurliana (Gates) Wiggins (Cactaceae) Mesembryanthemum crystallinum L. (Aizoaceae) Mesembryanthemum nodiflorum L. (Aizoaceae) Mirabilis laevis (Benth.) Curran var. crassifolia (Choisy) Spellenb. (Nyctaginaceae) Perityle incana A. Gray (Asteraceae) Phacelia floribunda Greene (Hydrophyllaceae) Plantago ovata Forssk. (Plantaginaceae) Rhus integrifolia (Nutt.) Brewer & S. Watson (Anacardiaceae) Spergularia macrotheca (Hornem.) Heynh. var. talinum (Greene) Jepson Sphaeralcea palmeri Rose (Malvaceae) Stephanomeria guadalupensis Brandegee (Asteraceae) Sphaeralcea palmeri, and Stephanomeria guada- lupensis. The succulent, perennial Cistanthe gua- dalupensis appears to be the most common of all of the species on the islet. It was in full flower during our visit in the early part of June, but other conspicuously flowering species included Baeriop- sis guadalupensis, Eschscholzia palmeri, Perityle incana, and two species of Hemizonia. The plant community of the central basin on the islet is dom- inated by Atriplex barclayana, Cistanthe guadalu- pensis, and Sphaeralcea palmeri with a strong pres- ence of Dudleya guadalupensis, Euphorbia misera, Lavatera lindsayi, Lycium californicum, Mammilla- ria blossfeldiana var. shurliana, Spergularia ma- crotheca, and Stephanomeria guadalupensis. On the upper slopes of the basin Baeriopsis guadalu- pensis, Coreopsis gigantea, Eschscholzia palmeri, Hemizonia greeneana ssp. greeneana, and H. pal- meri are prominent. The endemic Stephanomeria guadalupensis is most common on the lower slopes and bottom of the basin. The steep northern slope was the only location where Erysimum moranii, Phacelia floribunda and Rhus integrifolia were found. One individual plant of a putative interspe- cific hybrid between Hemizonia greeneana ssp. REBMAN ET AL.: FLORA OF TORO ISLET 147 FLoristic List OF TORO ISLET, BAJA CALIFORNIA, MExico. Note that Coll. # refers to the collection numbers Coll. # 6753, 6763 6758, 6882 6774 6765 6759 678la 6781b 6773 6757 6772 6755 6779 6761 6756, 6766 6775 6760 6770 6754, 6764 6780 6769 6778 6884 6784 6767 n/a Onn 7. 6776 6782 6783 6771 6762 6768 6883 greeneana and H. palmeri was found on the north- ern ridge. The flora of Toro Islet is very similar to that of Zapato Islet (also known as Islote de Afuera or Out- er Islet) with a few exceptions. It is noteworthy that Eriogonum zapatoense was not found on Toro, though an effort was made to look for it. This en- demic buckwheat species apparently only occurs on Zapato, where it is found on the upper ridge and basin rim and is relatively common there. Similar habitats were examined on Toro Islet, but it was not seen. Zapato Islet has a large basin that drops down to near sea level where plants such as Ap- hanisma blitoides occur. In general, Zapato appears to have a greater diversity of habitat types than Toro. Other species that occur on Zapato and not on Toro are Atriplex californica, Calystegia ma- crostegia ssp. macrostegia, Crassula connata, Crossosoma californica (which may no longer oc- cur on Zapato), Descurainea pinnata ssp. menziesii, Hutchinsia procumbens, Lavatera occidentalis, Oli- gomeris linifolia, Parietaria hespera var. hespera, Perityle emoryi, and Pholistoma racemosum. Plant taxa found on Toro Islet and not on Zapato include: Cryptantha maritima, Hemizonia palmeri, a puta- 148 tive Hemizonia hybrid, Hordeum murinum, Lupinus niveus, Mesembryanthemum nodiflorum, and Plan- tago ovata. All of these plants are found on the adjacent main island, which lies in closer proximity to Toro. While Toro Islet was explored quite fully, there are still possibilities for plants that were missed, particularly annual species. A botanical survey of Toro in early spring of a rainy year would probably identify additional annuals in the islet’s flora in gen- era such as Descurainea, Hutchinsia, Oligomeris, Parietaria, and Pholistoma, all of which have been recorded on Zapato Islet. MISCELLANEOUS BOTANICAL NOTES AND OBSERVATIONS FROM THE EXPEDITION In total, 149 plant collections were made during the entire expedition of which, four specimens doc- ument new distributional records for both native and exotic species not previously known to occur on the island. The new records of naturalized, ex- otic species include: Atriplex rosea L. (Rebman et al. 6817) and A. semibaccata R. Br. (Rebman 6750), both found near the airstrip in the center of the main island; and Schismus barbatus (L.) Thell. (Rebman 6752) found in the vicinity of the North- east Anchorage along the trail in Barracks Canyon. Of particular note was the discovery of a new native species for the island, Lonicera hispidula Douglas var. vacillans A. Gray. This pink-flowered honeysuckle is also found on Santa Cruz, Santa Catalina, and San Clemente islands of California, but is the first record (Rebman et al. 6800) of its occurrence in Mexico. Only one individual plant was found on the upper parts of a 5-meter-high rocky cliff cascade. Moran (1996) lists the weedy exotic Chamomilla suaveolens (Pursh) Rydb. (=Matricaria matrica- rioides (Less.) Porter) as part of the Guadalupe flo- ra, but it should be noted that the specimen (Wig- gins & Ernst 113) upon which this is based was misidentified and is actually the native species C. occidentalis (E. Greene) Rydb. Another plant spe- cies, Dichelostemma capitatum, was listed in Mo- ran (1996) as possibly introduced to Guadalupe Is- land. However, its presence on Toro and Zapato as well as on major portions of the main island does not support the concept that it was introduced. In preparation for the expedition, all of the data from specimens previously collected on Guadalupe Island and its adjacent islets that are housed in the SD Herbarium were compiled into a database. Dig- itized images, mostly scanned herbarium speci- mens, of the endemic plant taxa for the island ar- chipelago were also captured. Botanical specimens collected on the expedition were done so in replicate, whenever possible, so that specimens would be available for study in sev- eral herbaria. The first collections will be deposited at the SD Herbarium of the San Diego Natural His- MADRONO [Vol. 49 tory Museum, and duplicates will be in Mexican herbaria at HCIB in La Paz and BCMEX in Ense- nada. Plant materials including seeds and/or speci- men samples for genetic analyses were collected and sent to several institutions. Samples of species in the Asteraceae including Baeriopsis, Perityle in- cana, Stephanomeria, and the putative Hemizonia hybrid material were sent to the Jepson Herbarium (UC/JEPS) at the University of California Berkeley. Leaf material of Castilleja fruticosa was sent to the University of Washington (WTU) and samples of Lavatera lindsayi, Lupinus niveus, and Phacelia floribunda were sent to the University of South Da- kota. Triteleia guadalupensis material was sent to the University of Wisconsin for research on the Themidaceae. While conducting botanical explorations on Za- pato Islet during the expedition, some observations were made on the reproductive biology of two plant species endemic to the Guadalupe Island group. These observations are based only on morphology and their functionality still needs to be investigated further. It appears that Mammillaria blossfeldiana var. shurliana is a gynodioecious species with two sexual conditions. Although many of the cacti were not in flower at the time of our visit, a few flow- ering individuals showed evidence for two flower types occurring on different plants. Most flowering individuals appeared to have only bisexual flowers, with both functional stamens and pistils. However, a few plants were obviously different and seemed to be functionally female. These pistillate individ- uals had flowers with smaller tepals, abortive an- thers not producing pollen, and pistils with a larger size and increased number of stigmatic lobes. Plants showing these different floral characteristics grew in the same immediate vicinity on the islet. Gynodioecy is not uncommon in the Mammillaria species of Baja California since it has been docu- mented in M. dioica and some of its relatives in the region (Lindsay and Dawson 1952). However, this is the first report of this sexual condition in this cactus species. The other notable flower forms occurred in Cis- tanthe guadalupensis. Some individuals had flow- ers with “‘normal-sized’”’ petals (corolla diameter approximately 3—4 cm.) and yellow anthers, while other plants had “‘normal-sized’’ flowers but had pink-purple anthers. Furthermore, a fewer number of individual plants had significantly smaller flow- ers (corolla diameter approximately 2 cm.) and much shorter inflorescences. These plants with smaller flowers and inflorescences exhibited only yellow anthers and no fruit formation was seen. All of these three different flower forms were observed on individuals growing in close proximity to one another on Zapato. The comparable stages of flow- ering and similar moisture conditions of the local habitat seem to rule out any environmental cause for these floral variations. Therefore, it is hypoth- esized that Cistanthe guadalupensis is an andro- 2002] dioecious species. It appears that the individuals that have smaller flowers, shorter inflorescences, and no fruit formation are functionally staminate, while the plants with “‘normal-sized”’ flowers and viable pistils that develop into fruits are hermaph- rodites. The variability in anther color on different hermaphroditic plants is not yet understood and may not affect the reproductive system of this spe- cies. CONCLUSION The botanical data obtained from Toro Islet dur- ing this expedition not only helps to fill in the miss- ing pieces about the overall flora of Guadalupe Is- land and its adjacent islets, but it also yields im- portant information for conservation efforts. Due to the extinction of various plant species as a result of the ecological devastation caused by feral goats on the main island, the undisturbed Toro Islet is a very important resource for plant taxa that can be used for restoration of the main island if and when the goats are removed. The diversity of native and en- demic plants in this region is a rich heritage and serious conservation strategies need to be imple- mented in order to lessen the threats for their sur- vival into the future. REBMAN ET AL.: FLORA OF TORO ISLET 149 ACKNOWLEDGMENTS We are indebted to Drs. Michael Hager and Exequiel Ezcurra of the San Diego Natural History Museum and William Everett of the Endangered Species Recovery Council, for arranging and organizing the expedition. We thank the appropriate Mexican authorities for granting permits allowing the visitation to the island and the col- lection of plant species. We are grateful to the captain and crew of the Shogun for transportation to the island; Thom- as Thrailkill and Robert Gannon for field logistics; and Mel Cain for his remarkable piloting of the helicopter. We would also like to recognize Judy Gibson and botany vol- unteers at the SD Herbarium for their help in compiling a database of Guadalupe Island collections and scanning herbarium specimens. A special thanks to Patricia Beller for her efforts in obtaining the papers and permits needed to conduct the trip and for her help with the Spanish re- sumen. This material is based upon work supported by the National Science Foundation under Grant No. 0074462. LITERATURE CITED LINDSAY, G. AND E. Y. DAwson. 1952. Mammillarias of the islands off northwestern Baja California, Mexico. Cactus and Succulent Journal (U.S.A.) 24:77-84. Moran, R. V. 1996. The flora of Guadalupe Island, Mex- ico. Memoirs of the California Academy of Sciences 19. California Academy of Sciences, San Francisco, CA. REBMAN, J. 2001. The succulents of Islote Toro, Baja Cal- ifornia, Mexico. Cactaceas y Suculentas Mexicanas 46:52—55. MADRONO, Vol. 49, No. 3, pp. 150—157, 2002 GENETIC STRUCTURE OF SENECIO LAYNEAE (COMPOSITAL): A RARE PLANT OF THE CHAPARRAL GLENDA D. MArsH! California State University, Sacramento, 6000 J Street, Sacramento, CA, 95819 DEBRA R. AYRES Evolution and Ecology, One Shields Avenue, University of California, Davis, CA, 95616 ABSTRACT The genetic structure of Senecio layneae, a rare plant endemic to the fire adapted chaparral in the central foothills of the Sierra Nevada, California, USA, was investigated over the entire known range of the species. Genetic variation was assessed using 63 ISSR and 42 RAPD polymorphic DNA markers. Multivariate analysis differentiated populations from each of three counties and grouped two populations from El Dorado County together. ISSR markers more accurately portrayed population membership patterns than did RAPD markers. A species-wide F,; of 0.28 (by AMOVA) and a F.; of 0.22 within El Dorado County showed that significant genetic differentiation exists in the species and between local (within 5.5 km) populations. These results argue for maintenance of disturbed openings in the chapparal which this federally listed, threatened species can recolonise after fire, and for the use of local seed sources (within 5.5 km) for re-introduction efforts. Additionally, the preservation of multiple populations is indicated in order to maintain the existing pattern of genetic diversity across the landscape. Key words: ISSR, RAPD, genetic structure, S. layneae, chaparral Senecio layneae E. L. Greene (Composital), Layne’s Butterweed, is a perennial herb with an underground rootstock that forms small clones. It is restricted to open rocky areas within chaparral plant communities on gabbroic or serpentine-de- rived soils in the central Sierran foothill counties of Yuba, El Dorado, and Tuolomne in California (Cal- ifornia Native Plant Society 1994; USFWS 1996). Plants occur in sunny openings within the chaparral and die off as vegetation grows up around them (Baad and Hanna 1987). Flowering between April and June, the inflorescence is composed of disk flowers and a handful of unevenly distributed ray flowers. Primarily insect pollinated (G. Marsh per- sonal observation), its dandelion-like seeds are suit- able for dispersal by the wind. It is not known if a persistent, heat resistant seed bank of S. layneae exists in the soil beneath the chaparral. Senecio layneae was federally listed as threat- ened in 1996 (USFWS 1996). Chief threats include residential and commercial development, road maintenance, decreased fire frequency, off-road ve- hicle use, competition from invasive alien vegeta- tion, shading from native tree and shrub species, excessive horse grazing practices and gold mining (USFWS 1996). Populations of between 3 and per- haps 1000 plants (M. Baad personal communica- tion) are scattered within the chaparral of western El Dorado County including the Pine Hill gabbroic ' Present address: 2130 51st Street, Box 113, Sacra- mento, CA, 95817. E-mail: gdmarsh@juno.com. intrusion and adjacent serpentine (USFWS 1996). In Tuolumne County, several populations of be- tween 2 and 500 plants (A. Franklin personal com- munication) were first documented in 1984, and oc- cur in the chaparral dominated BLM Red Hills Management Area (BioSystems Analysis, Inc. 1984). In Yuba County two populations, one of ap- proximately 200 plants on public land, and the oth- er composed of scattered plants found on private lands, are found in chaparral and open Ponderosa Pine forest (Bureau of Land Management 1997). They were first reported in 1997, expanding the range of S. layneae northward 93 km. The protection of genetic diversity within species has become a priority for conservation efforts (Hol- singer and Gottleib 1991; Falk 1992). Such genetic diversity may be evident between individuals with- in a population or between the populations in a re- gion. The long-term objective is to maintain the evolutionary viability of the taxon, and maximize its chances for persistence in the face of changing environments (Huenneke 1991). Genetic diversity contributes to short-term ecological success in the face of stochastic environmental events such as lo- cal population destruction, pathogens, or herbivory (Holsinger and Gottlieb 1991). In rare plant pre- serve planning, an effort to preserve the species’ entire genetic diversity must be a goal (Frankel and Soulé 1981). The genetic variation maintained within a species is distributed among regions, populations, and in- dividuals within populations (Barrett and Kohn 2002] 1991). Variation in rates of gene flow, intensity of natural selection, and random genetic drift create a hierarchy which is referred to as genetic structure. Genetic structure arises when gene flow between populations is limited, favoring development of dis- tinct arrays of genetic characters within each pop- ulation (Wright 1951; Nei 1973). It is important to determine the spatial scale at which differentiation of populations occurs in order to understand the pattern of population divergence and microevolu- tion of a species. Preservation of this pattern should be a priority for species conservation (Driscoll 1998). Several life history factors, primarily those re- lated to pollen and seed dispersal, have been as- sociated with the development of genetic structure (Hamrick and Godt 1989). Wind carried seed, and an outcrossed breeding system, such as occurs in Senecio layneae (G. Marsh personal observation), promote gene flow, hampering the development of genetic structure and reducing the affects of genetic drift. Alternatively, Senecio layneae appears to re- colonize sunny openings left in the chapparal after disturbance such as fire or brush clearing, resulting in a historically patchy distribution. Small, patchily distributed populations would lead to the develop- ment of genetic structure through drift. Our primary research goal was to examine the genetic structure of this species and to infer the spatial scales at which gene flow is common and rare. Additionally, we will infer how historical fire regime could have influenced the genetic structure of populations seen today. Previously, much information about the genetic structure of plants was based almost entirely on data from electrophoretic surveys of soluble en- zymes (allozymes). Recently, RAPD-PCR-based methods have been used to investigate plant genetic Structure G-e., Huff et al. 1993: N’Goran et al. 1994; Baruffi et al. 1995; Ayres and Ryan 1997; Martin et al. 1997). These nuclear genetic markers are presumed to be selectively neutral and can re- veal patterns in variation due to neutral process such as random genetic drift and gene flow. This knowledge does not necessarily inform us regard- ing the arrangement of adaptively important traits across the landscape (Olfelt et al. 2001). However, even with these limitations, information from mo- lecular markers provides insight into population isolation due to restricted gene flow, and diver- gence due to drift which are potent factors in spe- cies evolution. A new type of marker, ISSR (inter-simple se- quence repeat), appears to be even more efficient than RAPD’s (random aplified polymorphic DNA’s) and has been used in recent investigations of nat- ural plant populations (Robinson et al. 1997; Wolfe et al. 1998a, b; Esselman et al. 1999). ISSR markers are repeat-anchored primers that amplify regions between SSR’s (simple sequence repeats or micro- satellites) (Tsumura et al. 1996). ISSR primers an- MARSH AND AYRES: GENETIC STRUCTURE OF SENECIO LAYNEAE 15] 1 Yuba County El Dorado County EN Tuolumne County uae 300 0 300 Kilometers Fic. 1. Locations of four sample sites for Senecio lay- neae in Yuba County, El Dorado County, and Tuolumne County, California, 1999. neal directly to SSR’s and no prior knowledge of the genome is required (Tsumura et al. 1996; Wolfe and Liston 1998). Additionally, ISSR’s produce a complexity of patterns as well as a level of poly- morphisms detected per single PCR experiment that largely exceeds that of RAPD’s (Zietkiewicz et al. 1994; Esselman et al. 1999; Ajibade et al. 2000). They are interpreted as dominant markers similar to RAPD data and are scored as binary data with ‘band present’ or ‘band absent’ (Wolfe et al. 1998a). Our secondary research goal was to com- pare the accuracy of ISSR and RAPD markers in determining genetic structure in S. layneae. MATERIALS AND METHODS Sample Collection Plant material was collected in the early spring when the dormant rootstocks began producing new growth. Four sites were sampled: Yuba County near Brownsville, Pine Hill and Cameron Park in El Do- rado County, and the Red Hills in Tuolumne Coun- ty (Fig. 1). One apical meristem was removed from 20 individuals from each of four populations. Plants were patchily distributed and sampled individuals were haphazardly chosen at least 2 meters apart to avoid sampling the same individual twice. Each sample was placed on ice until returned to the lab at which time they were stored at —70°C until DNA was extracted. DNA Extraction DNA was extracted according to Ayres and Ryan (1997) with the addition of a RNAase treatment between the two alcohol precipitations: 100 pl of TE (10 mM Tris-HCL and 1 mM EDTA, pH 7.8) and | unit of RNAase (Sigma) were added to the 152 MADRONO first DNA pellet and the tubes were then incubated at 35°C for 20 min. Precipitation in ethanol fol- lowed. The DNA was quantified from the absor- bance at 260 nm in a spectrophotometer. PCR and Electrophoresis Prior to screening primers, the optimum PCR an- nealing temperature was determined by running a temperature gradient reaction using a single acces- sion. The optimum annealing temperature was found to be 39°C for RAPD’s, and 54°C for ISSR’s. Amplification was performed with the following thermocycler programs: for RAPD’s, 94°C for 15 sec, 39°C for 30 sec, 72°C for 2.0 min, repeated 40 times, and then a 4°C holding temperature; for ISSR’s the program was identical except that the annealing temperature was 54°C. RAPD and ISSR PCR were performed in an Ep- pendorf Mastercycler gradient (Westbury, NY). DNA amplification reactions were performed in a volume of 15 pl containing approximately 30 ng of plant DNA, 0.20 pM/liter primer (Operon Tech- nologies, Inc., Alameda, CA, primers A4, Cl, C8, C10, D7, G8, G13, G14, G18, G19, H9 for RAPD); University of British Columbia kit 800, primers 807, 823, 836, 846, 848, 857, 859, 860 for ISSR), 200 wM/liter each of dATP cCTP, dGTP, dTTP (Promega, Madison, WI), 0.6 units Taq polymerase (Promega, Madison, WI), 3 wmol/liter MgCl2, 10% by volume MgCl2-free 10 reaction buffer A (Pro- mega, Madison, WI), and 10.5 wl of double dis- tilled water. The reaction mixture was overlain with a drop of mineral oil. PCR products were loaded on to 1.5% agarose gels for electrophoresis in 0.50 TBE buffer, followed by staining in ethidium bromide, and visualized and photographed under UV light. Molecular sizes of the visualized bands were estimated by referencing a 100 base-pair lad- der (Gibco Co.). Polymorphic, reproducible bands were scored as present or absent. Eighty-four decanucleotide RAPD primers from Operon Technologies, Inc. (Alameda, CA) and 84 ISSR primers from the University of British Co- lumbia were screened against one DNA sample to find primers that amplified DNA of S. layneae. The primers that produced bands were then subjected to a second evaluation using one accession from three populations (Yuba, Cameron Park, Pine Hill). Prim- ers that amplified DNA from all three populations were subjected to a third screening using three ac- cessions from each of the three populations (9 total DNA samples) to identify primers that produced polymorphic bands. Each of the above screenings was conducted two times to ensure reproducibility of results. Any primer which did not produce the same results was not used. Ultimately, 11 RAPD and 8 ISSR primers were selected for this study. [Vol. 49 Data Analysis ISSR and RAPD analyses Of 80 samples from which DNA was extracted, 74 accessions were ultimately included in all RAPD and ISSR data analyses. Five accessions with di- vergent band patterns were determined to be from a different Senecio species as at the time of sample tissue collection it was not possible to differentiate this second species of Senecio. These were dropped from further analyses. One accession was lost dur- ing DNA extraction. Fifty-eight RAPD and seventy-five ISSR poly- morphic markers were produced by PCR. Analysis showed that 28 markers had band patterns identical to other markers; these duplicates were dropped from further analysis. The remaining 42 RAPD and 63 ISSR markers were used in multivariate analyses of inter-individual and population genetic distance. For comparative data analyses of each marker type, a sub-sample of 35 markers of each type (for a total of 70) were randomly selected to provide an equal number of markers. Cluster analysis Matrices of RAPD and ISSR phenotypes, in which marker presence (1) or absence (0) was re- corded, were analyzed for inter-individual genetic distance separately and together using the Euclidean distance coeficient of the SIMINT subprogram of NTSYS-pc, version 2.01d (Rohlf 1993). Resulting genetic distance matrices were used to construct den- drograms using the SAHN subprogram and employ- ing unweighted pair group averaging (UPGMA) in NTSYS-pce. AMOVA Analysis of molecular variance (AMOVA) (Ex- coffier et al. 1992) was used to partition the genetic distance (assessed using Euclidean distance) among individuals within a population and among popula- tions. This hierarchical analysis of variance parti- tions the total variance into covariance components (Schneider et al. 2000). The covariance components are used to compute fixation indices in terms of in- breeding coefficients (Schneider et al. 2000). AMO- VA was originally developed for RFLP haplotypes, but has been used for RAPD phenotypes (Huff et al. 1993). By extension, it is also appropriate for ISSR phenotypes. The F,; statistic was computed and test- ed for significance after 1000 permutations (Schnei- der et al. 2000). These analyses were undertaken us- ing Arlequin ver. 2.0 (Schneider et al. 2000) avail- able at http:/Agb.unige.ch/arlequin/software/. Mantel test A Mantel test (Mantel 1967) of the correlation between genetic distances and geographic distanc- es, and the correlation between the RAPD and ISSR genetic distance matrices were calculated for all 2002] MARSH AND AYRES: GENETIC STRUCTURE OF SENECIO LAYNEAE 153 RAPD and ISSR Euclidean Distance PH CP PH CP PH — Pine Hill CP — Cameron Park T — Tuolumne Y - Yuba El Dorado Tuolumne Yuba eee OT ee Oo ee ee 2.83 3.82 4.81 5.80 Genetic Distance Fic. 2. UPGMA clustering of genetic distance of Senecio layneae individuals collected from Pine Hill (PH) and Cameron Park (CP) in El Dorado County, Tuolumne County (T), and Yuba County (Y). pairs of individuals using the MXCOMP program in NTSYS-pce (Rohlf 1993). The significance of the matrix was evaluated by comparing the observed Mantel test statistic Z, with its random distribution obtained after 1000 permutations. Interindividual geographical distances were found by calculating the Euclidean distance based on the UTM coordi- nates of each population. RESULTS Eleven RAPD primers and eight ISSR primers produced 42 and 63 distinct polymorphic bands, re- spectively. ISSR’s produced almost 2-fold more polymorphic bands per primer than RAPD’s (P(t = 4.125) > 0.001). Polymorphic bands ranged in size from 400 bp to 1800 bp. Genetic Structure of Senecio Laynea Multivariate analysis resulted in a dendrogram clearly showing three population clusters with all individuals grouped within their geographical pop- ulations at the county level (Fig. 2). While the El Dorado county populations of Pine Hill and Cam- eron Park were placed in a single large cluster, most individuals were grouped into generally discrete subclusters within the main county cluster (Fig. 2). Highly significant (P < 0.000001) genetic dif- ferences between populations were detected in the analysis of molecular variance (AMOVA) carried out on the genetic distance matrix. Of the total ge- netic diversity, 72% was attributable to individual differences within a population, and 28% to diver- gence among populations (Fy; = 0.28) (Table 1). Analysis of just the two El Dorado County popu- lations resulted in a F,, of 0.22. Analysis of Differences between RAPD and ISSR Equalizing the number of polymorphic bands for each marker type and analyzing each marker type separately resulted in somewhat similar aggrega- tions of individuals except for the Tuolumne Coun- ty population. This population was genetically dis- tinct when assayed by ISSR, but was intermixed with the Cameron Park and Pine Hill (El Dorado County) individuals when analyzed with RAPD markers (dendrogram not presented). The number of individuals placed in the wrong geographical population was significantly greater using RAPD’s than, ESSR’s. (0:01 <= P(y? = 9.407),.<.0.05)...An 154 TABLE 1. MADRONO [Vol. 49 AMOVA OF GENETIC DISTANCE IN SENECIO LAYNEAE. The total data set contained 74 individuals from four populations; El Dorado County analysis includes two populations from that county. P is the probability of obtaining a larger than observed variance component or fixation index by chance alone. Percentage Source of variation of variation Among all populations Dp Within populations 220i Among El Dorado County pop’s Dies) Within El Dorado County pop’s 67.29 AMOVA based on 35 ISSR markers resulted in a much stronger degree of between population dif- ferentiation than an AMOVA based on 35 RAPD markers (F,; = 0.38 and 0.26, respectively). Mantel Tests The correlation between genetic distance (as- sessed using all markers) and geographical distance separating individual plants was not significant (r = 0.277, P < 0.49). The correlation between ge- netic distance matrices based on 42 RAPD markers and 63 ISSR markers was significant, but low (r = 0.345, P < 0.002). DISCUSSION Significant genetic structure exists in S. layneae (Fsy = 0.28) resulting in three distinct geographic populations in El Dorado, Tuolumne, and Yuba Counties. Further, significant genetic structure (Fy, = 0.22) exists between two populations, separated by 5.5 km, in El Dorado County. These statistical findings were in agreement with multivariate clus- ter analysis. In contrast, in Wyethia reticulata, a clonal sunflower restricted to chaparral in El Do- rado County, populations separated by as little as 500 m had a distinct genetic identity (Ayres and Ryan 1997). The lack of significant correlation be- tween genetic distance and geographic distance in S. layneae could be interpreted in two ways: either gene flow is occurring among all populations and they are not isolated from each other, or populations have been isolated for a long enough time for in- dependent divergence to have occurred through random processes or natural selection. Given the preceding evidence, which supports substantial population divergence, we conclude that gene flow is extremely rare among populations of S. layneae, and populations have diverged through random ge- netic drift or selection. Several characteristics that may counter the de- velopment of genetic structure are known for S. layneae. Senecio layneae is primarily outcrossing with insect pollination (G. Marsh unpublished data and personal observation), disperses its seed on the wind, and has a perennial life history. Outcrossing generally reduces genetic structure and promotes higher genetic variation within populations. Wide seed dispersal prevents divergence among popula- Pp Fixation indices <0.001 FST = 0.279 <0.001 <0.001 ES =0216 <(0.001 tions via small amounts of long-distance migration and promotes higher genetic variation within pop- ulations. Perennial life history, especially long life, reduces the effects of drift and increases the chanc- es of migration, thus hindering divergence of pop- ulations and loss of genetic variation. (Loveless and Hamrick 1984; Hamrick and Godt 1989; Linhart and Grant 1996; Ayres and Ryan 1999). These fac- tors may be responsible for the intermingling of individuals from the Pine Hill and Cameron Park populations in El Dorado County within a single cluster: gene flow between the two sites, 5.5 km apart, does occasionally occur. Small populations of plants, scattered between these two populations, may act as a genetic bridge allowing this gene flow to occur. However, the species dispersal abilities do not extend to populations more than 90 km apart. The Red Hills Management Area sample site is ap- proximately 98 km south of the Cameron Park sam- ple site, and the Yuba County sample site is ap- proximately 93 km north of the Cameron Park site. A comprehensive search for additional populations acting as genetic bridges between these three areas would shed more light on the patterns of genetic diversity in S. layneae. Patchy spatial distribution can increase isolation and reduce gene flow, enhancing differentiation among populations if pollinator behavior reduces gene flow between patches. Patchy spatial distri- bution could also be at work in the apparent sub- structuring in the Pine Hill and Cameron Park pop- ulations. In addition, the ease and rapidity with which seeds of S. layneae germinate suggest that no seed bank exists, and so seed bank contributions probably do not play a role in the maintenance of within population diversity. Utility of ISSR Markers In this study, ISSR primers produced significant- ly more polymorphic bands per primer than did RAPD primers. More importantly, when equal numbers of polymorphic bands were used in cluster analysis of population membership patterns, ISSR- based genetic distance estimates more accurately portrayed population assignment of S. /ayneae in- dividuals than RAPD-based estimates. Wolfe et al. (1998a) found that UPGMA dendrograms derived from ISSR markers showed more highly resolved 2002] phylogenetic reconstructions of Penstemon popu- lations in the section Spectibilies than trees based on rDNA, cpDNA, and allozyme identity coeffi- cients. ISSR’s better resolved genotypes for three of four populations studied than did RAPD’s in ex- amining clonal diversity in rare Calamagrostis por- teri ssp. inseperata (Esselman et al. 1999). These results suggest a difference in the utility of RAPD and ISSR markers. In addition, the correlation be- tween RAPD-based and ISSR-based genetic dis- tance was low. These differences could arise due to differences in areas of the genome that each type of marker amplifies, because of higher error in RAPD markers, or because of higher precision in ISSR markers. Conservation Implications One of the main goals of conservation programs for species that are rare or threatened is to maintain existing levels of genetic variation (Avise 1994). While most of the genetic variation in Senecio lay- neae is found within populations, almost % of var- iation is found among populations. This is enough to argue for conservation of S. /Jayneae populations throughout its range. We also determined that pop- ulations separated by 5.5 km, while occasionally exchanging genes, are still quite distinct genetically (Fs; = 0.22), which argues for the use of local seed (within 5.5 km) for restoration projects. The nature of the disjunct and patchy distrubution of S. lay- neae populations is historical having to do with cy- cles of fire creating appropriate habitat within the chapparal. However, human activities, such as fire suppression and urban development, certainly influ- ence habitat availability in El] Dorado County ex- tinguishing local populations and further isolating remaining populations. Information about the fate and persistence of S. /ayneae individuals and pop- ulations throughout a fire cycle will be necessary to formulate a conservation strategy for this species that goes beyond management of local populations in each of the counties where it is currently found. Plants employ several mechanisms to cope with periodic fire. The responses to fire of three other rare plants found on the Pine Hill gabbro complex in El Dorado County demonstrate some of these strategies (D. Ayres unpublished data). Fire kills plants of the herbaceous trailing vine Calystegia stebbinsii but promotes seed germination from the soil seed bank. Plants grow rapidly and flower pro- fusely 2—3 yr after fire. As the canopy closes during the interfire period C. stebbinsii almost completely dies out, but populations can once again establish after fire as long as the soil seed bank has been replenished. The fire response of the low growing woody shrub Ceanothus roderickii is quite similar, except that the juvenile plants do not begin to flow- er until 5—6 years after fire, and some plants survive and flower under mature chaparral. Populations re- quire a fire-free period of at least 6 years to replen- MARSH AND AYRES: GENETIC STRUCTURE OF SENECIO LAYNEAE 155 ish the seed bank in order to exist in perpetuity. Fire leaves unharmed the underground rhizomes of W. reticulata and promotes vigorous flowering and seed set. This plant remains in the understory in the interfire period when the canopy closes over, but with much reduced flowering vigor. While we suspect that S. Jayneae does not have a fire resistant seed bank due to a relatively soft seed coat and an absence of any seed dormancy, this is but one of several questions regarding the fire response of S. layneae that needs to be ad- dressed in order to formulate a management strat- egy for this species. Other questions concern the survival of the caudex during the interfire period and during fires, and the environmental conditions that promote flowering and favor seedling estab- lishment. These biological constraints can be used to answer two key management questions; how long should the fire interval be, and how large should controlled burns be? If the caudex of S. Jayneae can survive both the canopy closure of the interfire period, and fire, we predict it will respond similarly to W. reticulata; abundant flowering shoots will emerge from the caudex, seed production will be increased several fold, and seedlings will establish in the fire’s ash. Under this scenario, precise fire intervals and areas are not critical management components for popu- lation survival as long as some areas burn some- time. However, if the caudex dies out during the interfire period or is killed by fire, and there is no seed bank, S. layneae will function as a fugitive species. It will require open patches of chaparral, near an existing reproducing population, to which the current crop of seeds can disperse and set seed before fire or canopy closure occurs. This is an en- tirely different fire survival strategy than the three other species described above and is similar to that of Furbish’s lousewort (Pedicularis furbishiae) (Menges 1990). The disturbance/successional niche for Furbish’s lousewort is defined by a river hy- drology in which patches of dense shrub thickets and trees are removed by scouring ice flows and bank slumping. The lousewort recolonizes the new- ly opened patch from water-born seeds. In this model, disturbance in the form of fire or ice flows opens up the regeneration niche (Grubb 1977) al- lowing recolonization by wind or water-dispersed seed from surviving populations. Populations die out as succession progresses, so species survival requires Ongoing disturbance. If this is the model for S. layneae, survival of the species will depend on both fire to create the regeneration niche and a supply of seeds to colonize the patch, either dis- persing from neighboring populations, or from sown seed collected from nearby populations. Fur- thermore, the genetic structure found in S. /ayneae supports this ‘fugitive’ model where loss of popu- lations due to catastrophic fires or lack of fires frag- ments populations and fosters genetic drift in the remaining populations. 156 Ideally, instead of focusing on the preservation of each extant population of each imperiled species as they occur today, the requirements for continued existence of all imperiled species will be incorpo- rated into an integrated management plan. This plan would include fire management, control of invasive exotic plants, protection of preserve boundaries, prevention of short-interval fires, and strategies to ensure not only species survival but maintain pat- terns of genetic diversity laid down through millen- nia of microevolution. 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Ge- nome fingerprinting by simple sequence repeat (SSR)-anchored polymerase chain reaction amplifi- cation. Genomics 20:176—-183. MADRONO, Vol. 49, No. 3, pp. 158-164, 2002 WOOD ANATOMY AND SUCCESSIVE CAMBIA IN SIMMONDSIA (SIMMONDSIACEAE): EVIDENCE FOR INCLUSION IN CARYOPHYLLALES S.L. SHERWIN CARLQUIST Santa Barbara Botanic Garden, 1212 Mission Canyon Road, Santa Barbara, CA 93105, U.S.A. ABSTRACT Simmondsia chinensis (Link) Schneider, endemic to dry areas of California, Arizona, and adjacent Mexico, is the sole species of Simmondsiaceae. Wood anatomy and cambial activity of this species are analyzed in view of the transfer of the taxon from Buxaceae or Euphorbiales to Caryophyllales s.l. The wood contains features considered primitive in dicotyledons: tracheids, diffuse parenchyma, and rays that are both multiseriate and uniseriate. These features are shared with Agdestidaceae, Rhabdodendraceae, and Stegnospermataceae, families considered basal to most of the Caryophyllales s.s. (“‘core Caryophyl- lales”’). Simmondsia has nonbordered perforation plates and successive cambia, features that occur widely in Caryophyllales s.1. Wood anatomy of Simmondsia is congruent with placement of the genus in Cary- ophyllales s.l. Simmondsia wood features reflect the desert habitat both quantitatively and qualitatively. Terminology and ontogenetic concepts of Simmondsia secondary xylem, successive cambial action, and periderm are contrasted with those in a previous study, and the problems of analysis of woods with cambial variants are discussed. INTRODUCTION Simmondsia chinensis, contrary to its species name, iS native to limited areas of arid hills and low mountain ranges of southern California, south- ern Arizona and adjacent portions of Mexico (Munz 1973). The genus has frequently been placed in Euphorbiales (see Goldberg 1986, Table I, for the treatments of Simmondsiaceae by 11 leading phylogenists). Simmondsiaceae has been placed in the family Buxaceae within the order by many workers. The unisexual flowers and tricoccoid fruits were suggestive of this relationship for many sys- tematists. However, Simmondsia was unique within Euphorbiales in having successive cambia. Succes- sive cambia have evolved in several groups inde- pendently (e.g., Menispermaceae, Gnetales); con- sequently, the presence of successive cambia was probably not considered a feature of prime taxo- nomic value. Recent phylogenies based on molecular data have opened possibilities in taxonomic interpreta- tion with regard to Simmondsia and some other genera with successive cambia. These newer views have effectively supplanted earlier thinking, so var- ious earlier phylogenies are not compared here. An expanded Caryophyllales (“‘caryophyllids”’) was proposed by Williams et al. (1994). Most notably, Droseraceae and Nepenthaceae were added, with Dilleniaceae the first branch on this clade. In sub- sequent phylogenetic constructions (Nandi et al. 1998; Soltis et al. 2000), families added to the ex- panded Caryophyllales include Ancistrocladaceae, Asteropeiaceae, Dioncophyllaceae, Frankeniaceae, and Tamaricaceae. Santalales are considered the outgroup for Caryophyllales s.l. In all of the phy- logenies utilizing molecular data, Buxaceae are not adjacent to Caryophyllales s.l., so the concept that Simmondsia belongs to Buxaceae is not supported. The present study is designed to compare wood and stem anatomy of Simmondsia to that of Caryophyl- lales s.1. that may be related. That purpose was also basic to the recent study of Rhabdodendraceae (Carlquist 2001a). Successive cambia occur in Car- yophyllales s.l. in Agdestidaceae, Amaranthaceae, Aizoaceae, Barbeuiaceae, Basellaceae, Caryophyl- laceae, Chenopodiaceae, Nyctaginaceae, Phytolac- caceae, Plumbaginaceae, Polygonaceae, and Steg- nospermataceae. The only families of Caryophyl- lales s.s. (‘core Caryophyllales’’) that lack succes- sive cambia are Cactaceae, Didieriaceae, and Portulacaceae. Thus, the presence of successive cambia in Simmondsia alone is a reason to compare Simmondsia to Caryophyllales. There have been previous accounts of Simmond- sia wood, notably those of Solereder (1885, re- ported as Brocchia), Van Tieghem (1897), Bailey (1980), and Carlquist (1982a). The present study goes beyond those studies by utilizing scanning electron microscopy (SEM) and by careful attention to ontogenetic phenomena (and associated termi- nology). Better understanding of successive cambia in Simmondsia will lead to a more accurate picture of successive cambia in dicotyledons and Gnetales. MATERIAL AND METHODS Stems of Simmondsia chinensis from a large shrub cultivated in the Santa Barbara Botanic Gar- den were fixed in 50% aqueous ethanol. Stems were taken in January and June, 2001, in order to compare degree of meristematic activity in the lat- 2002] CARLQUIST eral meristem. The stem of S. chinensis is hard enough to be sectioned, without softening, on a sliding microtome, but thin sections contain frac- tures. Some sliding microtome sections were stained with safranin and used for permanent slides. Some tangential sliding microtome sections were dried between clean glass slides, sputter coated with gold, and viewed with a Bausch & Lomb Nan- olab SEM. Portions of “‘bark’’ (tissue exterior to vascular bands) were fixed in 50% aqueous ethanol, softened for three days at 60°C in 8% ethylene di- amine, infiltrated, embedded with paraffin, and sec- tioned according to the method of Carlquist (1982b). These sections were stained with a safra- nin-fast green combination. Macerations were pre- pared with Jeffrey’s Fluid (equal parts of 10% chro- mic acid and 10% nitric acid) and stained with saf- ranin. The stems studied were between one and three cm in diameter. Vessel lumen diameter rather than outside vessel diameter was measured; for vessels oval in transec- tion, long and short chords were averaged. The ves- sel density recorded was based upon scans that did not include conjunctive tissue. If conjunctive tissue were to be included, the number of vessels per mm? would be about 50% lower. Both earlywood and latewood were included in the computation of quantitative vessel data. Terms are in accordance with the [AWA Committee on Nomenclature (1964) and Carlquist (2001b). The term “‘successive cam- bia’ follows the usage of Schenck (1893) and Pfeiffer (1926). RESULTS Secondary Xylem Growth rings inconspicuous, but evident on the basis of earlywood vessel diameter (Fig. 1); early- wood is usually not initiated at the beginning of each vascular band, but at some point within vas- cular bands. Vessels are virtually all solitary (Fig. 1); mean number of vessels per group, 1.04. Mean vessel lumen diameter, 21 fm. Mean number of vessels per mm’, 260 wm. Mean vessel element length, 163 wm. Mean vessel wall thickness, 2.8 wm. Perforation plates simple, nonbordered (Fig. 3, top). Helical thickenings present on vessel walls, some in the form of pairs of thickenings parallelling helices of pits (Fig. 3). Pit cavities of lateral wall vessel pits about 3 ym in diameter. Imperforate tra- cheary elements all tracheids, densely covered with fully bordered circular pits about 3 4m in diameter (Fig. 4). Mean tracheid wall thickness, 3.2 wm. Ax- ial parenchyma sparse, diffuse, composed of cells that are not subdivided. Rays uniseriate to wide multiseriate (Fig. 2); uniseriate rays are more com- mon than multiseriate rays. Most uniseriate rays are a single cell in height (and thus not easily seen in Fig. 2). Mean height of multiseriate rays, 212 pm. Mean width of uniseriate rays, 74 sm. Mean ray cell wall thickness, 1.5 im, walls lignified. Ray cell : SIMMONDSIA 159 wall pits simple. Ray cells predominantly procum- bent; square and upright cells relatively uncommon. Starch abundant in ray cells (Fig. 5). Secondary xylem nonstoried. Conjunctive Tissue and Cambial Action Conjunctive tissue is composed of bands five to ten cells thick radially (Fig. 1: tangential bands of thin-walled radial cells, containing phloem strands, intervening between tangential bands of dark sec- ondary xylem). As seen in radial section (Fig. 6), cells are mostly upright, a few square (and thus contrast with ray cells of the secondary xylem). Cell walls of conjunctive tissue are lignified and about 1.5 wm in thickness, but thin-walled nonlig- nified cells present on bark side of phloem strands (Fig. 7, above crushed phloem). Functional phloem cells, adaxial to the dense crushed phloem strand, are somewhat compressed due to sectioning rather than polygonal. What portion of the phloem cells that are not crushed are functional could not be de- termined. Continued production of secondary phlo- em is possible because earlier-formed phloem is progressively crushed. Solitary rhomboidal crystals are occasional in conjunctive tissue (Figs. 6, 8). Periclinal divisions can be seen in young conjunc- tive tissue (Fig. 9), but number of cell layers in the meristematic zone of the lateral meristem between the most recently initiated vascular cambium and the previous vascular band varies (e.g., Fig. 8). Vascular cambia (pointers at right edge, Figs. 7-9) actively produce secondary phloem and sec- ondary xylem: the secondary phloem occurs as iso- lated strands, whereas the secondary xylem mostly occurs as unbroken cylinders that extend around the stem. The first tracheids produced by a cambium are polygonal in outline, and their alignment in ra- dial rows is sometimes obscure because during maturation, the interfaces of the several faces shift. As divisions wane in each vascular cambium, a few thin-walled tracheids in radial rows are produced (Fig. 7, below phloem). Cambial activity produces phloem for a prolonged period, probably for several years because the abaxial portion of secondary phloem strands consist of numerous crushed phlo- em cells (Fig. 7, gray mass in center of photograph) and phloem cells produced by the cambium are in radial rows. Also, crushed phloem cells are always abaxial to apparently functional secondary phloem cells (Figs. 1, 7). Phellogen and Lateral Meristem Action The outer layers of the relatively mature stems studied here lack the cortical fiber strands reported by Bailey (1980) in the relatively young stems he studied. Such fiber strands had been shed from the stem I studied. Phellem, which consists of cells filled with dark-colored contents, is present on the stem surface. The phellem cells are narrower than cells in what is here termed diffuse lateral meristem 160 MADRONO [Vol. 49 ri DM Fics. 1-5. Stem sections of Simmondsia chinensis. 1. Transection (abaxial side above), showing alternation of bands of secondary xylem (each with associated phloem strands) and conjunctive tissue; vascular bands are not annual in extent; the pointers demarcate part of the earlywood of one year (above pointers) from part of the latewood of the preceding year (below pointers), several vascular bands are produced per year. 2. Tangential section of secondary xylem; a few large multiseriate rays are apparent (center), uniseriate rays are mostly inconspicuous because of their very small size. 3-5. SEM photographs from a tangential section of secondary xylem. 3. Portion of inner surface of a vessel, showing nonbordered perforation plate (top) and helical thickenings. 4. Outer surface of a tracheid, showing bordered nature and density of pits. 5. Starch grains in ray cells. Figs. 1, 2, magnification scale above Fig. 1 (divisions = 10 um; Figs. 3—5, scale bar in each figure = 5 wm). 2002] CARLQUIST: SIMMONDSIA 161 Fics. 6-9. Sections of stem of Simmondsia chinensis. 6. Radial section of conjunctive tissue to show cell shape and (center) rhomboidal crystal (secondary xylem at extreme left, secondary phloem at extreme right). 7. Strand of secondary phloem and surrounding tissues (conjunctive tissue above, secondary xylem below); crushed secondary phloem in abaxial part of phloem strand (pointer indicates site where vascular cambium was prior to cessation of active division. 8, 9. Transections of lateral meristem zone at periphery of stem; pointers at left in each indicate offset between outermost cells of the radial lateral meristem (radial files) and the innermost periderm cells; pointers at right indicate vascular cambium location. 8. Lateral meristem zone in which a vascular cambium has recently formed (no secondary xylem or phloem elements identifiable yet); rhomboidal crystal at bottom center. 9. Lateral meristem zone in which vascular cambium has yielded a vessel (extreme left) and some tracheids as well as some secondary phloem (right). Diagonal arrows denote recent divisions in the radial files of the lateral meristem zone; horizontal arrow (lower right) denotes a cell plate (obscured by cell contents adherent to it) that indicates a recent division in conjunctive tissue that is still somewhat meristematic. Figs. 6, 7, magnification scale above Fig. 6 (divisions = 10 pm). Figs. 8, 9, scale above Fig. 8 (divisions = 10 wm). 162 (it could also conceivably be termed secondary pa- renchyma). The diffuse lateral meristem cells are in radial rows; the term “‘diffuse’’ is used because di- visions do not occur synchronously in a single layer as in a vascular cambium, but are randomly dis- tributed throughout the meristematic zone (see Carlquist 1999a). A lateral meristem that consists of a single layer was observed in Barbeuia (Carl- quist 1999b) and also occurs in Nyctaginaceae (Carlquist unpublished data; data in earlier papers vary in interpretation and will be discussed in a later paper). More numerous divisions were ob- served in the stems collected in June than in those collected in January. The radial rows of lateral mer- istem cells are offset from the periderm which con- sists of a single layer of phellogen (narrow, like the phellem cells, but devoid of dark-colored com- pounds). In some places, there is a layer of paren- chyma between the phellogen and the lateral mer- istem files. No phelloderm cells were identified un- equivocally. The offset between the radial rows of the diffuse lateral meristem and the periderm is in- dicated by a pointer at the left in Fig. 8 and Fig. 9. The entirety of the periderm is illustrated in Figs. 8 and 9, which were selected to show primarily lateral meristem and vascular cambia. Origin of vascular cambia occurs within the ra- dial files of cells produced by the diffuse lateral meristem. Although only a small portion (for rea- sons of clarity) could be illustrated, study of the entirety of sections validates this interpretation. The origin of a vascular cambium (Fig. 8, pointer at right) can be distinguished from divisions of the lateral meristem because divisions of the vascular cambium form a single meristematic layer of divi- sions that are synchronous in tangential bands around the stem. Each vascular cambium soon pro- duces secondary xylem internally (adaxially) and secondary phloem abaxially (Fig. 9, pointer at left; vessels and a few tracheids in secondary xylem). The terminal products of a vascular cambium (Fig. 7) are described above. CONCLUSIONS Phylogenetic Position The occurrence of successive cambia is a char- acter widespread in Caryophyllales s.s. (“core Car- yophyllales’’) so its occurrence in families now added to an expanded Caryophyllales—Rhabdoden- draceae (Carlquist 2001a) and Simmondsiaceae is not surprising. ““Caryophyllales: s.s.’’ corresponds to the betalain-containing families plus Achatocar- paceae, Barbeuiaceae, and Molluginaceae, and the genera Limeum and Lophiocarpus (Clement et al. 1994). Simmondsia has characters generally consid- ered primitive in dicotyledons: presence of tra- cheids, presence of diffuse axial parenchyma, and presence of both multiseriate and uniseriate rays (Metcalfe and Chalk 1950: xlv, “fibres with dis- tinctly bordered pits; Kribs 1935, 1937). All of MADRONO [Vol. 49 these features are present in Rhabdodendraceae (Carlquist 2001a), now placed at the base of Car- yophyllales s.l., and in genera now placed at or near the base of Caryophyllales s.l. (Soltis et al. 2000). Simmondsia is placed by Hoot et al. (1999) and Soltis et al. (2000) near the base of Caryophyllales s.s. The other genera with the primitive features listed above include Agdestis (Carlquist 1999c) and Stegnosperma (Carlquist 1999a); Barbeuia has tra- cheids, but not the other character states mentioned above (Carlquist 1999b). One feature of possible ordinal significance is the presence of nonbordered perforation plates. These have been demonstrated in most Caryophyllales s.s. (see Carlquist 1999a, b, 2000). Nonbordered per- foration plates are newly reported here for Sim- mondsia (Fig. 3), and have recently been reported for some Caryophyllales s.l. such as Rhabdoden- draceae (Carlquist 2001a). Nonbordered perforation plates may be a symplesiomorphy in Caryophylla- les s.l. according to the above data and other ob- servations (Carlquist 2001b). Ecology Simmondsia is a desert shrub with only slight succulence in the leaves (Bailey 1980); not surpris- ingly, it has xeromorphic wood. The Mesomorphy Ratio (vessel diameter times vessel element length divided by vessel diameter) was reported to be 27.8 for Simmondsia by Carlquist and Hoekman (1985). A very similar value (24.4) can be derived from the present data if conjunctive tissue is not excluded. The similarity of the two reports is even closer if one notes that outside vessel diameter, rather than lumen diameter was used by Carlquist and Hoek- man (1985). Tracheids are conductively safe (excellent at con- fining embolisms to a single cell) compared to ves- sel elements. Fiber-tracheids and libriform fibers, by contrast, are nonconductive (see discussion in Carlquist 2001b). The presence of tracheids in Sim- mondsia is a feature of value in a xeromorphic hab- itat. Because of the presence of tracheids, vessel grouping in Simmondsia is virtually nil (1.04), in agreement with the correlation for dicotyledons as a whole claimed by Carlquist (1984). The value of tracheids in promoting conductive safety exceeds the value of vessel grouping (Carlquist 2001b). Ontogeny and Terminology The terminology in papers and books that deal with successive cambia is remarkably diverse, but more significantly, different interpretations often underlie the terms used. The present paper is not a proper venue for a review of this situation. How- ever, the paper by Bailey (1980) on Simmondsia is appropriate for comparison in view of the ontoge- netic interpretations as well as mature structures de- tailed in both the present paper and Bailey’s. In the present interpretation, a diffuse lateral 2002] meristem forms outside of the vascular cylinder, near the stem periphery. This lateral meristem pro- duces radial files of cells, producing parenchyma cells with primary walls, cells which remain rela- tively meristematic judging from recent divisions to be found in this region. Within the lateral meristem zone, a new vascular cambium is formed (usually while the preceding vascular cambium is still ac- tively producing secondary xylem and phloem). In Bailey’s (1980) interpretation, the zone I have termed lateral meristem is called conjunctive tissue (despite its lack of lignified secondary walls as found in conjunctive tissue in older parts of the stem). Bailey (1980) uses the term “‘extrafascicular cambium”’ for what I term the vascular cambium in each of the concentric vascular bands. Bailey (1980) claims that “‘the phellogen is ac- tually a region of transition where the peripheral conjunctive parenchyma of previous extrafascicular cambia undergoes further cellular subdivision; a true phellogen is lacking.” In the relatively young stems illustrated by Bailey (1980), phellogen might well be formed from cortical parenchyma as it is in many dicotyledons, but my studies indicate the ex- istence of a self-perpetuating phellogen, as so fre- quently described in dicotyledons. This phellogen is distinguished from the lateral meristem (outer- most conjunctive parenchyma of Bailey) not only by its tangentially narrower cell diameter but by an offset between the periderm files and the files of cells in the radial parenchyma (Figs. 8, 9). These two differences would be difficult to explain if peri- derm were ontogenetically continuous with the files of cells of the lateral meristem. The vascular cambia produce strands of second- ary phloem externally and cylinders of secondary xylem internally. In my interpretation, quite ordi- nary rays are produced by each cambium. In Bai- ley’s (1980) interpretation, “‘conjunctive tissue ini- tials produce raylike structures of conjunctive tis- sue; true vascular rays are absent.’’ This interpre- tation has not, to the best of my knowledge, been offered in any genera with successive cambia other than in Bailey’s study of Simmondsia. The differences in interpretation detailed above show that careful analyses of successive cambia and other cambial variants still need to be under- taken. The diversity of interpretations and terms for the anatomical phenomena is still considerable. The number of different interpretations and terms within instances of successive cambium occurrence seems unlikely to be matched by an equal diversity of on- togenetic mechanisms. Rather, cellular arrange- ments have been viewed differently by different workers. In part, the diversity of interpretations and terminology may derive from microtechnical con- siderations. Thin sections such as can be cut with a rotary microtome, are desirable for revealing cell lineages and histological details clearly. The hard- ness of many stems with successive cambia has, on the contrary, led to preparation of sliding micro- CARLQUIST: SIMMONDSIA 163 tome sections in which soft tissues do not section well or are too thick for cell development sequenc- es to be revealed clearly. The use of rotary micro- tome sections of material that has been chemically softened to a suitable degree seems the best solu- tion to this dilemma. Embedding in resin or plastic is an alternative microtechnical possibility. LITERATURE CITED BaILey, D. C. 1980. Anomalous growth and vegetative anatomy of Simmondsia chinensis. American Journal of Botany 67:147-161. CARLQUIST, S. 1982a. Wood anatomy of Buxaceae: cor- relations with ecology and phylogeny. Flora 172:463— 491. . 1982b. The use of ethylene diamine in softening hard plant structures for paraffin sectioning. Stain Technology 57:311-—317. . 1984. Vessel grouping in dicotyledon woods: sig- nificance and relationship to imperforate tracheary el- ements. Aliso 10:505—525. . 1999a. Wood and stem anatomy of Stegnosperma (Caryophyllales): phylogenetic relationships; mature lateral meristems and successive cambial activity. In- ternational Association of Wood Anatomy Journal 20: 149-163. . 1999b. Wood anatomy, stem anatomy, and cam- bial activity of Barbeuia (Caryophyllales). Interna- tional Association of Wood Anatomy Journal 20:43 1— 440. . 1999c. Wood anatomy of Agdestis (Caryophyl- lales): systematic position and nature of the succes- sive cambia. Aliso 18:35—43. . 2000. Wood and stem anatomy of phytolaccoid and rivinoid Phytolaccaceae (Caryophyllales): ecol- ogy, systematics, nature of successive cambia. Aliso 19:13-29. . 2001a. Wood and stem anatomy of Rhabdoden- draceae is consistent with placement in Caryophyl- lales sensu lato. International Association of Wood Anatomy Journal 22:171—181. . 2001b. Comparative wood anatomy, 2nd ed. Springer Verlag, Berlin, Germany. AND D. A. HOEKMAN. 1985. Ecological wood anat- omy of the woody southern California flora. Inter- national Association of Wood Anatomy Bulletin, new series, 6:319—347. CLEMENT, J. S., T. J. MABRY, H. WYLER, AND A. S. DREID- ING. 1994. Chemical review and evolutionary signif- icance of the betalains. Pp. 247—261 in H.-D. Behnke and T. J. Mabry (eds.), Caryophyllales. Evolution and systematics. Springer Verlag, Berlin, Germany. GOLDBERG, A. 1986. Classification, evolution, and phylog- eny of the families of dicotyledons. Smithsonian Con- tributions to Botany 58:1—314. Hoot, S., S. MAGALLON, AND P. R. CRANE. 1999. Phylog- eny of basal eudicots based on three molecular data sets: atpB, rbcL, and 18S nuclear ribosomal DNA se- quences. Annals of the Missouri Botanical Garden 86: 1-32. IAWA COMMITTEE ON NOMENCLATURE. 1964. Multilingual glossary of terms used in wood anatomy. Konkordia, Winterthur, Switzerland. Kriss, D. A. 1935. Salient lines of specialization in the wood rays of dicotyledons. Botanical Gazette 96:547— oy 164 MADRONO . 1937. Salient lines of specialization in the wood parenchyma of dicotyledons. Bulletin of the Torrey Botanical Club 64:177—186. METCALFE, C. R. AND L. CHALK. 1950. Anatomy of the dicotyledons. Clarendon Press, Oxford, U.K. Munz, P. A. 1973. California flora and supplement. Uni- versity of California Press, Berkeley, CA. Nanpt, O. I., M. W. CHASE, AND P. K. ENprRgssS. 1998. A combined cladistic analysis of angiosperms using rbcL and nonmolecular data sets. Annals of the Missouri Botanical Garden 85:137—212. PFEIFFER, H. 1926. Das abnorme Dickenwachstum. Handb. der Pflanz. 9(2):1—272. Gebriider Borntraeger, Berlin, Germany. SCHENCK, H. 1893. Beitrage zur biologie und anatomie der Lianen. II. Schimpers Botanischen Mittheilen der Tro- pischen 5:1—271. [Vol. 49 SOLEREDER, H. 1885. Uber den systematischen Wert der Holzstruktur bei den Dicotyledonen. R. Oldenbourg, Miinchen, Germany. SOLTIs, D. L., P. S. So_tis, M. W. CHASE, M. E. Moegrt, D. C. ALBACH, M. ZANIS, V. SAVOLAINEN, W. H. HAmN, S. B. Hoot, M. EF Fay, M. AXTELL, S. M. SWENSEN, L. M. PRINCE, W. J. KRESS, K. C. NIXON, AND J. S. FARRIS. 2000. Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Botanical Journal of the Linnean Society 133:381—461. VAN TIEGHEM, P. 1897. Sur les buxacées. Annales des Sci- ences Naturelles, Botanique, series 8, 5:289—301. WILLIAMS, S. E., V. A. ALBERT, AND M. W. CHASE. 1994. Relationships of Droseraceae: a cladistic analysis of the rbcL sequence and morphological data. American Journal of Botany 81:1027—1037. MAbRONO, Vol. 49, No. 3, pp. 165-168, 2002 A NEW SUBSPECIES OF NAVARRETIA LEUCOCEPHALA (POLEMONIACEAE) FROM VERNAL POOLS IN EASTERN WASHINGTON CURTIS R. BJORK Marion Ownbey Herbarium, Washington State University, Pullman, WA 99163 cbjork @ wsu.edu ABSTRACT Navarretia leucocephala Benth. subsp. diffusa is newly described from numerous populations in a distinctive vernal pool landscape in the Channeled Scablands of Eastern Washington. Its affinities appear closer to subspecies of N. leucocephala in the California floristic province, than subspecies minima, which is the only other member of the N. leucocephala group known to grow within the range of the Flora of the Pacific Northwest (Hitchcock and Cronquist 1973). The range of subspecies diffusa closely corre- sponds with the central of three channels of the glacial Spokane Flood events and is segregated from populations of subspecies minima by expansive Palouse loess uplands, where vernal pools are absent. Just as no plants of subspecies diffusa were found outside this central flood channel, no plants of sub- species minima were found within the range of subsp. diffusa. Plants of N. leucocephala subsp. diffusa are easily distinguished from subspecies minima in Washington by the fewer flowers within the much more openly diffuse heads, and by the conspicuously puberulent calices, fewer seeds per capsule, longer corolla lobes and higher filament insertion. It is distinguished from other subspecies by the consistently puberulent calices, low seed number and the relatively simple outer inflorescence bracts. Key words: Navarretia, Polemoniaceae, vernal pools, Columbia Plateau, Channeled Scablands The Navarretia leucocephala Benth. complex in- cludes four species in North America: N. fossalis Moran, N. leucocephala Bentham, N. myersii PS. Allen & A. G. Day (two subspecies) and N. pros- trata (A. Gray) E. Greene, and one in South Amer- ica: N. involucrata Ruiz & Pavé6n (Day 1993a). Na- varretia leucocephala is defined as having five sub- species: leucocephala, bakeri (H. Mason) A. G. Day, minima (Nuttall) A. G. Day, pauciflora (H. Mason) A. G. Day, and plieantha (H. Mason) A. G. Day (Day 1993a). Two of the subspecies are widespread; subspecies /eucocephala occurs in the Great Valley of California and in southwestern Oregon (Day 1993b), and subspecies minima is found from California to north-central Washington, western Idaho and to Utah. During a study of vernal pools of the Columbia Plateau, populations of Na- varretia leucocephala encountered in one subre- gion of the Columbia Plateau vernal pool province stood out as being highly distinctive, leading to the morphological comparison with the other subspe- cies of N. leucocephala presented here. TAXONOMY Navarretia leucocephala Benth. subsp. diffusa Bjork subsp. nov.—TYPE: USA, Washington State, Lincoln County, Swanson Lakes Wildlife Management Area. Floors of vernal pools on scabland basalt flows. 680 m. T24N R34E S4 NEA/4. June 18, 1997 Curtis R. Bjérk 3229 (Ho- lotype, WS, Isotype WTU). Planta annua, erecta vel decumbens, 0.5—4.7 cm lata; caulis centralis 1.0—13.4 cm alta, ad medium 0.3—0.7 mm crassus; capitula hemispherica, (1) 4— 10 (15) mm lata, diffusa, plerumque ramificans, flo- ribus (1) 5—20 (30); bracteae exteriorae lobis (0) 2— 6 (8); calyx pilis brevis crispis plerumque recur- vatis, lobi plerumque integri; corolla inclusa, alba, lobi lineares; stamina in sinibus lobarum corollum inserta, inclusa, stigma superantibus; semina (1) 2 G6): Plants annual, erect to decumbent, 0.5—4.7 cm wide; central stem 1.0—13.4 cm high, 0.6 + 0.1 mm thick at midlength; heads hemispheric, (1) 4—10 (15) mm wide, diffuse and generally branching, with (1) 5—20 (30) flowers; outer bract lobes (0) 2— 6 (8); calyx conspicuously puberulent with crisped, mostly recurved hairs, lobes mostly entire; corolla included, white, lobes linear; stamens included, in- serted near corolla-lobe sinuses, surpassing the stig- ma; seeds (1) 2 (3). Paratypes. U.S.A. Washington, Lincoln County: adjacent to Knack Road, 3.5 miles SSW of Telford, T25N R35E S29 NE % of NW %, Bjork 3228 (WS), 18 June 1997; Swanson Lakes Wildlife Manage- ment Area, 1 mile West of Florence Lake, June, 1999, Bjork s.n. (WS); Swanson Lakes Wildlife Management Area, 2 miles East of refuge head- quarters, T25N R34E S35 NE %, 19 June 1997, Bjork 3250 (WS); 3.4 miles SSE of Swanson Lakes, along Seven Springs Dairy Road, 13 June, 1998, Mark Fishbein 3439 (WS); Large vernal pool along Swanson Schoolhouse Road, T25N R34E S31 NW % of NW %, Bjork 6161 (WS). Comparative morphology. Plants of this distinc- tive group of populations are here named Navar- retia leucocephala subspecies diffusa and are clear- 166 MADRONO ly distinguished from all other subspecies of N. leu- cocephala by their consistently and conspicuously puberulent calyx, their generally much more diffuse heads and their relatively simple outer bracts (Table 1). It is placed within N. leucocephala rather than with any other species or in its own specific rank due to the similarity of plant architecture and the cymose inflorescence to those of subspecies N. leu- cocephala (in contrast especially to N. myersii and N. prostrata) (Crampton 1954), and due to the low seed number (in contrast with N. fossalis). Addi- tionally, it differs from the South American species N. involucrata by flower color and the simpler leaves and bracts. The subspecies of N. leucoce- phala with the greatest morphological similarity to subsp. diffusa appears to be subsp. pauciflora of Northern California. Both have few seeds per cap- sule (usually 2 in both subspecies), few flowers per head, high filament insertion and corollas with nar- row throats and linear lobes. Specimens of subspe- cies diffusa are distinguished from those of subspe- cies pauciflora by the often broader (to 15 mm, though still few-flowered) and more diffuse heads, the consistently white flowers (versus the generally bluish flowers of subspecies pauciflora), the sim- pler outer bracts and the taller stature (central stem length to 10 cm). Also similar to subspecies diffusa is subspecies bakeri of Northern California and Southwestern Oregon. The fewer flowers per head and fewer seeds per capsule distinguish subsp. dif- fusa from subsp. bakeri. Subspecies diffusa differs from subsp. leucocephala in the included corollas and the higher filament insertion. Subspecies dif- fusa is distinguished from subsp. plieantha by the white versus bluish flowers, fewer flowers per head and by the lower seed number. Subspecies diffusa differs from subsp. minima by the simpler bracts, fewer flowers per head, lower seed number, lesser central stem length and width, narrower head width, higher filament insertion, ca- lyx length and the fewer calyx lobe divisions (Table 2). Plants of subspecies diffusa are also distin- guished from subsp. minima by the nearly simul- taneous maturation of flowers within a head. Unlike in subsp. diffusa, there are typically some flowers in subsp. minima heads that bear mature seeds while others are not yet in anthesis. The heads of subsp. diffusa are openly branching, so much so that the calyx bases are often clearly seen within. The calyx lobes of subsp. diffusa are unequal and longer in relation to the tube, ranging from 0.8—1.5 x the tube length, in contrast to those of subsp. minima, which are usually subequal in length and 0.5-1.0 X the tube length. Additionally, one to three of the ribs leading to the lobes in subsp. dif- fusa maintain their width and herbaceous tissue to the base of the calyx tube. The ribs are often wider than the intervening membranes, which is appar- ently unique within section Navarretia (Day 1993b) and the N. leucocephala complex. The calyx tube in subsp. diffusa is conspicuously puberulent MORPHOLOGICAL COMPARISONS OF THE SUBSPECIES OF NAVARRETIA LEUCOCEPHALA AND N. L. SUBSP. DIFFUSA SuBSP. Nov. Outlier values are shown in parentheses. Data are in part from Day 1993b, with additional observations from herbarium specimens (WS, OSC). TABLE 1. diffusa plieantha pauciflora bakeri 2-10 13-24 30—60 minima 2-11 (4) 7-16 (20) leucocephala 1-16 (10) (1) 4-10 (15) (1) 5-20 (30) (O) 2-6 (8) 1-3 15-20 20-80 2-22 9-36 15—80 Central stem length (cm) Head width (mm) Flowers per head Outer bract lobes 6-12 6-30 4-12 0-10 1.7-3.5X 0.8-3.0 gen. + equal none 20-80 6-10 4-8 9-16 8—30 1.2—-2.5X 1.0-2.3X 4—10 (O) 4—16 (30) A-14 (0) 4-19 0-6 1.0—2.2x 1.5—3.0X gen. + equal gen. none dense white Bract lobe 2° divisions 1.0—2.0x 1.5-3.0X 1.0—2.0X gen. unequal (1) 2-3 1.5-3.0X 0.6-3.0X gen. + equal gen. none gen. sparse white Bract length/head width ratio Calyx tube/lobe length ratio gen. + equal none Calyx lobe relative lengths Calyx lobes divided Calyx pubescence Corolla color gen. sparse gen. bluish + linear incl. gen. sparse gen. bluish + linear incl. gen. sparse gen. white + jinear incl. gen. sparse white + linear incl. + oblong incl. + ovate excl. Corolla lobe shape Corollas included/exserted Filament insertion Seed number at sinuses at sinuses in throat at sinuses at sinuses in throat 1—2 (3) (2) 3-5 (6) 2—4 2-3 AF) [Vol. 49 2002] BJORK: wa | A) yy awh WGA on. \\ > i Xi OK: A es i pe a. AG og Wyo 7 4 Ad) WY Sls yim Bia. 1. NAVARRETIA LEUCOCEPHALA SUBSP. DIFFUSA 167 Calamagrostis breweri was effectively lectotypified by the citation of Brewer 2128 as the type (Hitchcock and Chase 1950). We have examined these specimens. Brewer 2128 and Lemmon s.n. are individuals of the north- ern populations, with relatively wide leaves that rolled up as they wilted. Only Bolander 6098 has Tioga Pass C. muiriana Mt. Dana Carson Pass Mt. Eddy Mt. Jefferson Mt. Hood 0.40 0.60 0.80 1.00 Hedrick’s (1971) genetic similarity Fic. 2. Cluster diagram based on Hedrick’s similarities of Calamagrostis breweri (Mt. Hood, Mt. Jefferson, Carson Pass, and Mt. Eddy populations) and C. muiriana (Tioga Pass and Mt. Dana populations), calculated from isozyme pattern frequencies. truly setaceously involute leaves and represents the southern form. Kearney (1898) apparently observed the same morphological variation within C. breweri that prompted this study. He split the taxon into two species, C. breweri as the name was commonly ap- plied, and a new species, C. Jemmoni, characterized by broader leaves. He stated that C. lemmoni was “intermediate between C. deschampsioides and C. breweri”’ (Kearney 1898). He cited Lemmon s.n. as the type specimen. Lemmon s.n. is a broad-leaved northern plant, different from the majority of plants to which the name C. breweri has been traditionally applied. However, both Lemmon s.n. and Brewer 2128 are broad-leaved northern plants. The name C. lemmoni was appropriately relegated to synon- ymy with C. breweri by Hitchcock (1912). Kear- ney’s attempt to split C. breweri left the northern form with two names, and the narrow-leaved south- ern plant nameless. Doubtless this confusion re- sulted from the fact that leaves of both Lemmon s.n. and Brewer 2128 had wilted and rolled up tightly before pressing, so that they superficially ap- pear to be setaceously involute. Nygren (1954) detailed the cytology of C. brew- eri and rediscovered the its morphological varia- tion, but did not treat it taxonomically. He found that plants of two southern populations (near Tioga Pass in Mono County and Mt. Dana in Tuolumne County) were tetraploids (2n = 28), and plants from a northern population (near Carson Pass, Al- pine county) were hexaploids (2n = 42). Nygren also noted that the tetraploid and hexaploid forms could be distinguished by the shape of “‘the upper- most leaf of the straw [culm]? (Nygren 1954). More recently, Greene (1993) acknowledged the range of morphology, describing C. breweri leaves as “‘flat or involute.”’ CONCLUSIONS Kearney (1898) was correct that Calamagrostis breweri as traditionally understood includes two entities that differ in morphology and habitat. Per- haps the differences between the two taxa are due to consistent differences in chromosome number; ploidy level in itself can affect the morphological traits and habitat preference like those that differ- entiate the two forms of C. breweri (Tal 1980). Ploidy levels can represent barriers to gene flow, 174 and therefore populations that differ in chromo- some number may be treated as separate species (Harlan and de Wet 1971). However, ploidy level may mean little taxonomically; grasses of two ploi- dy levels may mingle in a population without ob- vious morphological or ecological differentiation (Hultquist et al. 1997; Keeler et al. 1987). Taxon- omists should avoid encumbering the taxonomic lit- erature with species that differ only in ploidy level (Lewis 1980), but in C. breweri the southern (pre- sumably tetraploid) and northern (presumably hexaploid) forms exhibit sufficient morphological, ecological, leaf anatomical, and isozyme differenc- es that they can reasonably be recognized as dif- ferent species. We name the southern species after John Muir, pioneering naturalist and first president of the Sierra Club. Muir wrote extensively about the Sierras and in particular the Yosemite area. In the summer of 1869, he spent more than a month in Big Tuolumne Meadows “sketching, botanizing, and climbing among the surrounding mountains” (Muir 1894, p. 70). He must frequently have walked or slept on this small reedgrass that carpets the floodplain meadows of the Yosemite region. Calamagrostis muiriana B. L. Wilson and Sami Gray sp. nov.—TYPE: USA: California: Tuol- umne County: Yosemite National Park; Dana Fork, Tuolumne River, Elevation: 9525 feet. 1 August 1997. B. L. Wilson and S. Gray 8909. (Holotype: OSC; isotypes, CAS, NY, RSA, UC, US, UTC). Gramen pusillum alpinum, maxime similare Cal- amagrostidi breweri, sed tetraploideum et foliis an- gustioribus trinerviis. Gramen perenne caespito- sum, 12—34 cm altum, culmis florentibus folia ex- cedentibus. Folia perangusta glauca, vaginis folior- um marginibus imbricatis, ligulatis 0.8—2.2 mm longis, laminis filiformibus glaucis glabris, 4—12 cm longis, 0.2—0.4 mm latis, venis ternis. Inflores- centia paniculata erecta, pauciflora, atroviolacea, 1.9-7.5 cm longa, ramis expansis. Spiculae atro- violaceae, uniflorae, rachilla sterili pilis albis, glum- is aequalibus, 3—4.5 mm longis, atroviolaceis, at- tenuatis vel minutae aristatis. Lemma hyalinum vel atroviolaceum, 2.5—4.0 mm longa, apicale denti- culatum, dentibus quatuor, pusillis, atroviolaceis, pilis calli albis 0.3—0.6 mm longis, arista lemmatis geniculata, 3.3-6.0 mm longa, infra medium cari- nae orienti. Palea hyalina, lemmati aequanti. An- therae maturae 0.9—2.5 mm longae, purpuratae. Species tetraploidea, chromosomata 28. Gramen abundans in pratis alpinis apricis, dis- tributum in montibus excelsis Californiae cen- tralis. A Calamagrostide breweri laminis angustioribus venis ternis, paniculis pusillioribus, pilis calli bre- vioribus distinguenda. Plants cespitose, with intravaginal shoots. Young plants densely tufted. Old plants spreading outward MADRONO [Vol. 49 while dying in the middle and thus forming rings 3 dm or more in width. Foliage glaucous, 6-15 cm long. Leaf sheaths open, glabrous to retrorsely short-pubescent. Ligules 0.8—2.2 mm long, entire, rounded apically. Leaf blades involute, well-devel- oped leaves 4—12 cm long 0.2—4 mm wide as in- volute, (but earliest leaves of innovations with blades ca. 0.5 cm long and 0.2—0.4 mm wide), leaf blades abaxially scabrous on veins and glabrous or scabrous between them, adaxially pubescent; leaf blades of innovations with 3 (—5) veins and usually 7 sclerenchyma bundles; leaf blades of culms with 5 veins. Leaf tip straight-sided, not prow-shaped. Flowering culms taller than the foliage and 12—35 cm tall. Inflorescence a spreading panicle, few- flowered, 1.9—7.5 cm long and 0.4—3.0 cm wide, dark purple or rarely straw-colored. Spikelets one- flowered, with a sterile rachilla about half as long as the floret and covered with long white hairs. Glumes equal in length, 3—4.5 mm long, thin-tex- tured, purple, often with hyaline margins distally (occasionally straw-colored), acute to apiculate, mi- nutely awned, or attenuate, rounded on the back or keeled distally, glabrous (occasionally scabrous), sometimes with hairs on the keel. Lemma 2.5—4.0 mm long, thin-textured, hyaline or dark purple, gla- brous or scabrous, usually with purple on veins dis- tally, the veins extending as four short (ca. 0.5 mm long) teeth. Callus beard hairs white, 0.3—0.6 mm long. Lemma awn purple, arising from below the middle of the lemma, 3.3—6.0 mm long, geniculate, extending beyond the glumes. Palea hyaline, pig- mented along veins distally, about as long as the lemma. Mature anthers 0.9—2.5 mm long, purple. 2n = 28. Descriptions of Calamagrostis breweri are, in most instances, descriptions of Calamagrostis mui- riana. We therefore re-describe Calamagrostis breweri sensu stricto here: Plants cespitose, with intravaginal (occasionally extravaginal) shoots. Young plants densely tufted. Old plants spreading outward while dying in the middle and thus forming rings ca. 1.5 dm in di- ameter Foliage glaucous, 10—20 cm long. Leaf sheaths open, scabrous. Ligules 1.7—4.1 mm long, entire to erose. Leaf blades flat but readily rolling when dry, well-developed leaves (2—) 10-15 cm long, 0.4—0.6 mm wide when rolled, the innovation leaves 0.9—1.1 mm wide when flat, (earliest leaves of innovations only slightly reduced), the culm leaves 1.3—1.7 mm wide when flat, leaf blades abaxially scabrous, adaxially pubescent; leaf blades of innovations with 7—9 veins and 9-11 abaxial sclerenchyma bundles; leaf blades of culms often with 11 or more veins. Leaf tip prow-shaped. Flow- ering culms taller than the foliage and 29-54 cm tall. Inflorescence a spreading panicle, few-flow- ered, 5.7—8.4 cm long, and 0.7 to 5.2 cm wide, pale to dark purple. Spikelets one-flowered, with a ster- 2002] Fic. 3. Calamagrostis muiriana. A. habit. B & C. spike- lets. D. floret. E. cross section of leaf blade. FE cross sec- tion of leaf sheath. ile rachilla about half as long as the floret and cov- ered with long white hairs. Glumes subequal in length, the lower 3.1—4.9 mm long; the upper 3.3— 4.5 mm long. Glumes thin-textured, pale to dark purple or sometimes greenish, often with hyaline margins distally (occasionally straw-colored), acute to apiculate, minutely awned, or attenuate, rounded on the back or keeled distally, glabrous (occasion- ally scabrous), sometimes with hairs on the keel. Lemma 2.6—4.0 mm long, thin-textured, hyaline or dark purple, glabrous or scabrous, usually with pur- ple on veins distally, the veins extending as four short teeth. Callus beard hairs white, 0.3—1.2 mm long. Lemma awn purple, arising from below the middle of the lemma, 3.4—5.5 mm long, geniculate, extending beyond the glumes. Palea hyaline, pig- mented along veins distally, about as long as the lemma. Mature anthers 1.3—2.6 mm long, purple. 2n = 42. WILSON AND GRAY: CALAMAGROSTIS 175 Calamagrostis breweri Calamagrostis muiriana Fic. 4. Distribution of Calamagrostis breweri (@) and C. muiriana (*) in Oregon and California. KEY TO ALPINE REEDGRASSES OF WESTERN NORTH AMERICA 1. Leaves extremely thin and involute, 0.25—0.4 mm wide as rolled, with 3 (—4) veins, the tip straight- sided; panicle length 1.9—5.7 cm; callus beard hair length 0.3—0.6 mm; range south of Sonora Pass (Yosemite area and south; Fig. 4) ..... C. muiriana 1. Leaves narrow, flat in life but readily rolling when wilted, 0.4—0.6 mm wide as rolled, with 7 or more veins, the tip prow-shaped; panicle length 5.7—8.5 cm; callus beard hair length 0.3—1.2 mm; range north of the Sonora Pass (Carson Pass area north to Mt= Hoods @Oreson)i 24 5...... 2: C. breweri s. str. ACKNOWLEDGMENTS We thank Dr. Aaron Liston for his guidance, and Susan Nugent for initiating this study. Heather Laub and Mike Roantree collected plants at Mt. Hood and Mt. Jefferson, respectively. We thank Dr. Richard Halse, curator of the Oregon State University Herbarium, for assistance in ob- taining herbarium specimens. Jan Van Wagtendonk and Leslie Chow provided the permit necessary to collect in Yosemite National Park. The study was supported finan- cially by the Mt. Hood National Forest and the Native Plant Society of Oregon. We thank Dr. Craig Greene for his helpful comments and for making available a copy of his dissertation on Calamagrostis. Finally, we thank the following institutions for loans of herbarium specimen: 176 MADRONO Rancho Santa Ana Botanical Garden, Humboldt State University, California Academy of Science, and the Uni- versity of California at Berkeley. LITERATURE CITED ABACUS CONCEPTS, INC. 1988. StatView 512+, Version 1.2. Berkeley, CA. ABRAMS, L. 1940. Illustrated flora of the Pacific states; Washington, Oregon, and California, Vol. 1; Ophiog- lossaceae to Aristolochiaceae: ferns to birthworts. Stanford University Press, Menlo Park, CA. ANONYMOUS. 1995. Rare, threatened, and endangered plants and animals of Oregon. Oregon Natural Heri- tage Program, Portland, OR. CHuNG, M. G., J. L. HAmrick, S. B. JONES, AND G. S. DERDA. 1991. Isozyme variation within and among populations of Hosta (Liliaceae) in Korea. Systematic Botany 16:667—684. GREENE, C. W. 1993. Calamagrostis. Pp. 1243-1246 in J. C. Hickman (ed.), The Jepson manual: higher plants of California. University of California Press, Berke- leven Orne HARLAN, J. EF AND J. M. DE WET. 1971. Toward a rational classification of cultivated plants. Taxon 20:509—517. Hircucock, A. S. 1912. Gramineae Pp 82-189. In W. L. Jepson (ed.), A flora of California: Part II. Cunning- ham, Curtis, and Welch, San Francisco, CA. AND A. CHASE. 1950. Manual of grasses of the United States. USDA Miscellaneous Publication No. 200. Government Printing Office, Washington, DC. HuLtToqulistT, S. J., K. PR. VoGEL, D. J. LEE, K. ARUMUGAN- ATHAN, AND S. KAEPPLER. 1997. DNA content and Chloroplast DNA polymorphisms among switch- grasses from remnant Midwestern prairies. Crop Sci- ences 37:595—598. KEARNEY, T. H. 1898. A revision of the North American species of Calamagrostis. U.S. Department of Agri- culture Division of Agrostology Bulletin 11:16. KEELER, K. H., B. KWANKIN, P. BARNES, AND D. W. GAL- BRAITH. 1987. Polyploid polymorphism in Andropo- gon gerardii. Genome 29:374—379 Lewis, W. H. 1980. 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Watson (ed.), Geological survey of California:botany, Vol. II. John Wilson & Son University Press, Cambridge, MA. WENDEL, J. E AND N. E WEEDEN. 1989. Visualization and interpretation of plant isozymes. Pp. 5—45 in D. E. [Vol. 49 Soltis and P. S. Soltis (eds.), Isozymes in plant biol- ogy. Dioscorides Press, Portland, OR. WHIPPLE, J. 1981. A flora of Mount Eddy, Klamath Moun- tains, California. Master’s thesis, Humboldt State University, Arcata, CA. YEH, E C., R.-C. YANG, AND T. BOYLE. 1997. Popgene version 1.21: Microsoft Windows-based freeware for population genetic analysis. Department of Renew- able Resources, University of Alberta, Edmonton, Al- berta, Canada. APPENDIX 1 SPECIMENS EXAMINED * = specimen included in morophometric study; ** = type specimen Calamagrostis muiriana: USA: California: Fresno Co., Baxter Lake, 6 Sep 1959, DeDecker 1119 (CAS); N of Kings River, Red Mt. Basin, 16 Aug 1961, Hardham 8748 (CAS); Silver Pass in Fish Creek Country; Sierra Nat’! Forest; 19 Sep 1912, Hatton H-109 (CAS); Death Pond, Kaiser Ridge, 12 Aug 1928, Jepson 13229* (UC); Bear Cr. Watershed, Mt. Hilgard, 20 Aug 1951, Quibell 680 (RSA); Bear Cr. Watershed, 2 mi W of Mt. Hilgard, 21 Aug 1951, Quibell 767 (RSA); Bighorn Lake, head- waters N Fk Mono Cr., 2 mi W of Red & White Mt., 31 Aug 1952, Quibell 1575 (RSA); Bighorn Lake, headwa- ters N Fk Mono Cr., 2 mi W of Red & White Mt., 31 Aug 1952, Quibell 1583 (CAS); Bear Creek, near Florence Lake, 7 Sep 1954, Quibell 4985 (OSC, RSA); Upper French Canyon Basin (near Bishop), 6 Aug 1955, Quibell 5387 (RSA); Colby Meadows, 24 Jul 1952, Raven 4677 (CAS), Laurel Creek, 13 August 1953, Raven 6174 (CAS); Rose Lake, 8 Aug 1954, Raven 7858 (CAS); Bench Lake, 25 Jul 1956, Raven 9846 (CAS); Humphrey’s Basin, W of Mt. Humphreys, 12 Aug 1937, Sharsmith 3165* (CAS, UC); Mono Pass, 118°50’W, 37°25'N, 10 Sep 1959, Thomas 8144 (CAS); Second Recess, 18 Jul 1953, Raven 5694 (CAS); Inyo Co., Paiute Pass; 11,300— 11,409 ft., 22 Jul 1934, Ferris 8877* (CAS, UC); Rock Cr. Lake Basin, Mosquito Flat, 14 Jul 1946, Howell 22277 (CAS); Mono Mesa, 26 Jul 1946, Howell 22730 (CAS); Big Pine Lakes, 5th Lake, 5 Aug 1947, Howell 23767 (CAS, RSA); Rock Cr. Lake Basin (Long Lake), 22 Jul 1931, Peirson 9398 (RSA); Rock Cr. Lake Basin, 17 Jul 1931, Peirson 9399 (RSA); Rock Cr. Lake Basin, Ruby Falls, 20 Jul 1934, Peirson 12617* (CAS, RSA, UC); Coyote Ridge, 7 Aug 1950, Raven & Stebbins 217 (CAS); Coyote Ridge, 2 Aug 1950, Raven & Stebbins 227* (UC); Mono Pass; Ruby Lake, 16 Jul 1936, Robinson 689 (RSA); Lone Pine Canyon, E of Mt. Muir, 20 Aug 1937, Sharsmith 3310* (UC); Mono Mesa near junction of Mono, Inyo, & Fresno Co., 37°28'N, 118°46'W, 9 Sep 1961, Thomas 9747 (CAS); Madera Co., Lyell Fork of Merced River, Yosemite Nat’] Park, 1 Aug 1931, Blasdale s.n. (UC); Minarets, 23 Aug 1918, Grant 1585 (CAS); Minarets, 23 Aug 1918, Grant 1587 (CAS); Mariposa Co., Soda Springs by the Tuolumne, 15 Aug 1894, Cong- don s.n. (CAS); Yosemite Nat’! Park, % mi N of Tenaya Lake, 18 Jul 1951, Stebbins 5005* (UC); Emeric Lake, T2S R24E S8, 15 Sep 1935, Thomas 490* (UC); May Lake, 15 Jul 1977, Vale s.n. (CAS); Mono Co., Slate Cr. Basin, E of Mt. Conness, 7 Sep 1934, Clausen 922 (CAS); Tioga Pass, Yosemite Nat’! Park boundary, 16 Aug 1936, Ferris 9824* (CAS, UC); Tioga Pass, just S of entrance to Yosemite Nat’] Park, Heller 15461 (CAS); Mono Pass, 16 Aug 1944, Howell 20631 (CAS); Saddlebag Lake, payee 2002] head of Lee Vining Cr., 18 Jul 1968, Howell & True 4432 (CAS); Saddlebag Lake, head of Lee Vining Cr. 18 Jul 1968, Howell & True 4435 (CAS): Slate Cr. Basin, 26 Aug 1937, Keck 4591 (CAS, UC); HM Hall Natural Area, Green Lake, N base of Mt. Conness, 27 Aug 1927, Keck 4610 (CAS); HM Hall Natural Area; saddle between Mt. Conness & East Plateau, 27 Aug 1927, Keck 463] (CAS); Slate Cr. Valley, Inyo Nat'l Forest, 26 Jul 1954, Krucke- berg 3602 (CAS, RSA); Slate Cr., 17 Aug 1954, Munz 19987 (RSA); Slate Cr., Aug 1954, Munz 20088 (RSA); Saddlebag Lake, 18 Jul 1968, Rose 68/51] (CAS, HSC, RSA); Tioga Pass, 14 Aug 1930, Stanford 2029 (RSA): summit of Dana Plateau; 10 km W of Mono Lake, 19 Jul 1951, Stebbins 5006* (UC): Mono Basin, Parker Cr. Ba- sin, | Aug 1987, Taylor 9210* (RSA, UC); Mono Pass, 4th Recess, 37°26'N, 118°48’W, 11,000—12,000 ft., 13 Sep 1954, Thomas & Thomas 4630 (CAS): Mt. Conness Re- gion, NE of Steelhead Lake, 11 Sep 1959, Twisselmann 5700 (CAS); Saddlebag Lake, 16 Aug 1933, Wolf 5334 (RSA); below Highway 120 near Yosemite E entrance, 11 Aug 1970, Wood 242 (HSC); near Tioga Pass, TIN R25E $30, 22 Aug 1936, Yates 6301 (RSA, UC); Tioga Pass, TIN R25E S30, 22 Aug 1936, Yates 6304* (CAS, UC); a mile above Conness Lake, Convict Cr. drainage, 26 Aug 1968, Zufeldt 2 (CAS); Tulare Co., Kern Lake, 11,000— 12,100 ft. CAP 2074* (UC); Mt. Whitney, 11 Jul 1910, Clemens s.n. (CAS); Yosemite Nat’] Park, Mt. Conness, 19 August 1897, Dudley 1703 (CAS); between Reflection Lake & Harrison Pass, 8 Aug 1940, Howell 16048 (CAS); Little Five Lakes Basin, 29 Jul 1942, Howell 17379 (CAS); Sky Parlor Meadow, Chagoopa Plateau, 1 Aug 1942, Howell 17520 (CAS): Center Basin, 26 Jul 1948, Howell 25042 (CAS, RSA); Rock Creek, 21 Jul 1949, Howell 25568* (CAS, UC); Mineral King vicinity, Fare- well Gap, 20 Jul 1951, Howell 28013 (CAS, RSA); Min- eral King Vicinity, White Sheif Region, 21 July 1951, Howell 28088 (CAS); Diamond Mesa, 25 Jul 1954, Kehr- lein (CAS); Kern Nat'l Forest (Mt. Whitney District), Morrow 5 (CAS); Mineral King Game Refuge, Sequoia Nat'l Forest, N of Franklin Lakes, 8 Aug 1966, Rice 5/6 (OSC, RSA): Kern Plateau, Twisselmann et al. 11280 (RSA); Chicken Spring Lake, Kern Plateau, 31 Jul 1970, Twisselmann et al. 17396 (CAS, RSA): Tuolumne Co., Tuolumne Meadows, September 1866, (Bolander?) 6098* (UC); Tuolumne Meadows, Yosemite Nat’! Park, Babcock 3627 (UC); Slide Canyon W of Finger Peak near N boundary of Yosemite Nat’! Park, 31 July 1934, Barthol- omew s.n. (UC); Mt. Lyell Quadrangle, 2 miles W of Mt. Gibbs, 6 Aug 1937, Bufford s.n.* (UC); Yosemite Nat’l Park, Lake Ireland, 29 Jul 1917, Clemens s.n. (CAS); Mt. Dana, 16 Aug 1894, Congdon s.n.* (CAS, UC): Yosemite Nat’! Park, Mt. Conness, 9 Aug 1934, DeRoy 115 (CAS); Gaylor Lake (upper), Yosemite Nat’l Park, Hall 11864 (CAS, UC); Tioga Pass, Yosemite Nat’! Park, 13 Sep 1922, Hall s.n. (CAS); Dog Lake, 6 Aug 1944, Howell 20034 (CAS); Gaylor Lake, 12 Aug 1944, Howell 20372* (CAS, UC); Lyell Fk of Tuolumne River, 25 Jul 1936, Lee 2334* (UC): Yosemite Nat’! Park, 12 Mar 1909, Lemmon 1897 (CAS); Tioga Pass, 30 Aug 1957, Rose 57127 (CAS, RSA, UC); Lyell Fk. of Tuolumne River, upper Lyell Can- WILSON AND GRAY: CALAMAGROSTIS ee yon, 25 Jul 1933, Sharsmith 189 (CAS, UC); Dana Mead- ows, west base of Mt. Dana, 15 Aug 1933, Sharsmith 414* (UC); Mt. Dana, west slope, | Aug 1931, Sharsmith S00 (UC); Mt. Dana, NW plateau, 10 Sep 1934, Sharsmith 2100 (CAS, US); Mt. Dana, NW slope, 2 Aug 1933, Sharsmith 257B (UC); Gaylor Lake, Yosemite Nat’! Park, 23 June 1976 Vale & Wagnon s.n. (CAS); Tuolumne Meadows, Mt. Lyell Quadrangle, 22 Aug 1936, Yates 6325* (CAS, RSA, UC): Tuolumne Meadows, Yosemite Nat’! Park, 8500—9500 ft., Jul 1902, Hall & Babcock 3627 (UC). Calamagrostis breweri: USA: California: Alpine (?) Co., near summit of Carson’s Pass, Brewer 2/28** (GH, US); Lake Winnemucca ca. 2 miles south of Carson Pass, 17 Aug 1996, Gray & Wilson 8270 (OSC); Amador Co., Woods Lake, summit of Carson Pass, 20 Jul 1951, Steb- bins 5009* (UC); Eldorado Co., Echo Lake, 11 Aug 1981, Best s.n.* (CAS); E side Wright’s Lake, T12N RI6E, 16 Jul 1977, Stebbins 7771* (CAS); Trail from Suzie to Heather Lake, 17 Aug 1928, Wolf 3338* (RSA); Nevada Co., Basin Peak, 27 Aug 1968, True 4590* (CAS); 1 mi SW of Mt. Lola, White Rock Lake, 7920— 8400 ft., 28 Jul 1976, Trowbridge SOO0* (CAS); 1 mi SW of Mt. Lola, White Rock Lake, 28 Jul 1976, Trowbridge S074 (CAS); Placer Co., Donner Pass, 10 Aug 1903, Heller 7130* (CAS, OSC, UC); Siskiyou Co., English Peak, Diamond Lake, 2 Aug 1968, Oettinger 345* (RSA); English Peak, Marble Mt. Wilderness Area, Shasta Ridge, 3 Sep 1969, Oettinger 1613* (RSA, UC); Shasta Ridge, N slope of English Peak, 3 Sep 1969, Oettinger 1613 (HSC); S side Caribou Lake, T37N RIOW S35, 19 Aug 1980, Renner 2506 (HSC); above Sugar Lake, T40N ROW S31, 4 Aug 1969, Sawyer 1803* (OSC); above Sugar Lake, 4 Aug 1969, Sawyer 1803 (HSC); W of Little Duck Lake, T40N ROW S19, 14 Sep 1972, Smith & Sawyer 5813 (HSC); Mt. Eddy, north face, T40N R5W S7, 25 Jul 1976, Whipple 1576 (HSC); Mt. Eddy, north slopes, T40N RSW S7, 20 Jul 1977, Whipple 2006 (HSC): near Little Crater Lake, T40ON R5W S5&6, 5 Aug 1977, Whipple 2183 (HSC); 1.45 mile by road north of Parks Creek Sum- mit, T41N R6W S33, 16 Aug 1996, Wilson & Gray 8258 (OSC); Trinity Co., Stuart Fk. drainage, west of Morris Lake, T36N RIOW S22, 24 Aug 1997, Ferlatte 1899* (HSC, UC); Lower Canyon Cr. Lake, 10 mi W of Dedrick, Salmon-Trinity Alps, 11 Jul 1939, Hitchcock & Martins 5413* (CAS, UC); Echo Lake, T35N ROW S3, 12 Aug 1994, Rolle 853 (OSC); California, no county, [no lo- cation], Lemmon s.n.** (US); [no location], 18 Feb 1909, Lemmon s.n. (US); Oregon: Hood River Co., Mt. Hood (SE; upper Sahalie Falls), 2 Aug 1924, Henderson 1045 of 1924* (CAS, OSC); Mount Hood Meadows Ski Area, T3S R9E S4, 11 Sep 1995, Laub s.n. (OSC); Mt. Hood, Heather Canyon, T2S R9E S24, 1 Aug 1999, Nugent s.n. (OSC); Mt. Hood, T2S R9E S34, Poff s.n. (OSC); Mt. Hood, T2S R8.5E S13, 4 Sep 1979, Siddall s.n.* (OSC); Paradise Park, Mt. Hood, 11 August 1926, Thompson 1660, Linn Co., Jefferson Park, 16 Aug 1946, Prescott s.n., Marion Co., Mt. Jefferson, TIOS R5E S11, 9 Sep 1992, Roantree, s.n.* (OSC); Mt. Jefferson Wilderness Area, SE endof Russel Lake, TIOS R8E S11, 18 Oct 1995, Roantree, s.n. (OSC). MADRONO, Vol. 49, No. 3, pp. 178-185, 2002 PLANT ZONATION IN A SHASTA COUNTY SALT SPRING SUPPORTING THE ONLY KNOWN POPULATION OF PUCCINELLIA HOWELLII (POACEAE) LARRY LEVINE P.O. Box 4783, Arcata, CA 95518 1-levine @northcoast.com MARY BACCA H. T. Harvey & Associates, 3150 Almaden Expressway, Suite 145, San Jose, CA mbacca @harveyecology.com K. O. FULGHAM Department of Rangeland Resources and Wildland Soils, Humboldt State University, Arcata, CA 95521-8299 fulghamk @axe.humboldt.edu ABSTRACT Three small salt springs adjacent to state highway 299 west of Redding, California, support the only known population of the grass, Puccinellia howellii J.1. Davis. The common halophyte grass, Distichlis spicata (L.) E. Greene, and Puccinellia each dominate different areas within the springs. In 1991, a highway realignment encroached on the Puccinellia, and in partial mitigation, an attempt was made to convert a Distichlis-occupied area to Puccinellia. The subsequent re-occupation by Distichlis forced a reconsideration of the restoration rationale and methods, and raised concern for the potential of Distichlis to replace Puccinellia elsewhere. But a TWINSPAN of systematically chosen samples suggested that the two grasses are members of distinct vegetation types associated with different hydrology. Salinity and growth monitoring of stands of Puccinellia and Distichlis suggested that the former tends to occupy areas continually irrigated by spring discharge, while the latter tends to occupy areas that are less directly irrigated, where salt can accumulate during the warm months, but also where precipitation during the cool months can lower the salinity enough to temporarily permit the growth of glycophytic annuals. Triglochin maritima L. and Juncus bufonius L. were also monitored. Greenhouse tests showed that Puc- cinellia is adversely affected by the high salinity typical of the Distichlis areas during summer. The association of P. howellii with continual surface flow should be considered when selecting and preparing revegetation sites, and when surveying for new populations. Key words: Puccinellia howellii, salt spring, salt marsh, Distichlis spicata, vegetation zonation, halophyte, endemic Adjacent to State Route 299, approximately 32 km west of Redding, California, near the juncture with Crystal Creek Road, and within the boundaries of the Whiskeytown Recreational Area, are three mineral springs totaling 0.49 hectares (Fig. 1). The low grass cover and the bare and rocky areas con- trast sharply with the adjacent canyon live oak and blue oak woodland. The springs discharge from nu- merous points, producing a sheet flow over much of the site as the water moves downhill. Salinity ranges from 15—35 dS/m (mmho/cm), approximate- ly half that of sea water, with a calcium content of 1—1.8 g/liter. Discharge volume and chemistry re- main stable year round (CH2M HILL 1991-1992). Though initially alkaline (pH 9—9.6) the water acid- ifies (pH 7-5) as it flows away from the discharge points (Fulgham et al. 1997), presumably due to the influence of decomposing organic matter. Typical for this region, precipitation is restricted to a period from mid-fall to mid-spring. Two halophytic perennial grass species dominate the site, usually in separate stands, Puccinellia how- ellii J.1. Davis, ““‘Howell’s alkali grass,”’ a cool sea- son species endemic to this site, and Distichlis spi- cata (L.) E. Greene (including D. stricta (Torr.) Rydb.), “‘salt grass,’ a warm season species wide- spread in both coastal and inland settings in North America. Puccinellia howellii was first recognized as a dis- tinct species in 1990 (Davis). The closest species morphologically is P. pumila (Vasey) A. Hitchc., found in coastal marshes from Washington to Alas- ka, rarely south to California (Kartesz 1999). Al- though commonly referred to as “alkali grass,” most Puccinellia are associated with saline habitat of neutral pH. Although Puccinellia howellii is in- cluded on the California Native Plant Society List 1B (“‘rare, threatened or endangered in California and elsewhere’’), it currently has no state or federal legal status (CNPS 2001). Puccinellia howellii stands range from sparse to dense, and consist either of individual tufts or a 2002] OREGON CALIFORNIA WHISKEYTOWN ae LAKE RED BLUFF PACIFIC ° OCEAN -% © 10 20 30 40 50 MILES Fic. 1. Location of the study area west of Redding, Shas- ta County (Bacca 1995). dense, turf-like growth. Plants at maturity can range in stature from 2.5 to 20 cm or taller, excluding the length of the inflorescences. The tuft form reaches the greatest height. The structure of the tufts sug- gests that individuals rarely persist longer than sev- eral years under field conditions. Triglochin mari- tima L. often co-occurs with Puccinellia in the wet- ter locations. While Puccinellia tends to occupy the central portions of the springs, Distichlis is often found at the periphery, adjacent to the surrounding non-sa- line vegetation. In winter and spring, when Distich- lis is dormant, the areas occupied by Distichlis of- ten support a sparse to dense cover of the annual grass species found outside the springs. At this lo- cation, Distichlis appears to spread primarily via rhizomes and stolons; little seed production has been observed. In contrast, Puccinellia reproduces primarily from seed. Juncus bufonius L., a glycophytic (non-halo- phytic) annual, seasonally occupies some otherwise barren areas within the springs. The exotic and po- tentially invasive Atriplex rosea L. is present, but was uncommon during the data collection portion of this study. In 1991, a realignment of State Route 299 en- croached on the salt springs. The California De- partment of Transportation (CalTrans) salvaged Puccinellia tufts from the construction zone and transplanted them to an adjacent 176 m? area re- claimed from a deposit of roadside spoil occupied by Distichlis. An aerial photo predating the spoil suggested that the area had previously been occu- pied by Puccinellia. The site was prepared by re- moving approximately % m of soil, which was found to be dense with Distichlis rhizomes. After transplanting, the Puccinellia tufts quickly became established, and at first produced considerable re- LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING NT) cruitment from seed. However, by the second year, Distichlis rhizomes and stolons began to re-enter from the periphery. By the third year, Puccinellia tufts were dying and not being replaced by seed- lings, and by 1998, Distichlis had replaced Pucci- nellia in most of the transplant area. This unantic- ipated outcome raised questions about the potential for Distichlis to replace Puccinellia elsewhere. Concurrent with the transplant project, CalTrans funded an ecological study of P. howellii, which was conducted by the authors between 1993-1995. Drawn from that study (Fulgham et al. 1997), this paper describes plant zonation within the springs, and provides information about the relative habitat requirements of the component species. METHODS Sampling. In May 1993, the visible extent of the three springs was systematically surveyed for veg- etation analysis. Parallel transects were laid out 3.05 m apart. Along each transect, an initial sam- pling position was selected randomly, and subse- quent positions spaced at 3.05 m intervals. Species percent cover was estimated using a tripod-mount- ed optical point-intercept bar (Cover-Point/ESCO) with 20 sighting positions along its 1 m length. At each sampling position, the bar was placed perpen- dicular to the transect, randomly either left or right. Each bar setting constituted a sample. The combi- nation of systematic and randomized elements in the sampling design ensured a uniform coverage, while providing a considerable degree of indepen- dence between samples. Because the Distichlis growth for the current year had not yet appeared, it was surveyed by using the dead growth from the previous year, which at the time was still structur- ally intact. In addition to the collection of species data, the surface moisture at each intercept point was re- corded in one of four categories, depending on whether it appeared dry, moist, saturated (i.e., glistening’), or with standing water. Vegetation analysis, association. The cover sam- ples were analyzed with TWINSPAN (Hill et al. 1988). Of the 551 samples, only the 394 with =5% cover were used. Pseudospecies cut levels were set at 5, 15, 33, and 67, and each given equal weight. Cover from species found in five or fewer samples were combined into a category “‘misc’’. Vegetation analysis, moisture. A subset of 235 vegetation samples was used to compare the sur- face moisture associated with each species. As de- fined, these were plots with =15% total cover, and where one species contributed >50%. In reality, the large majority of the samples that met these criteria had one species exceeding 66%. To compare the relative moisture between samples, the percent cov- ers for the moisture categories were weighted and summed to form a quasi-continuous moisture in- dex. ““Dry”’ cover was weighted by 0.25, “‘moist’’ 180 MADRONO [Vol. 49 TABLE 1. TWINSPAN VEGETATION GRouPS, MEAN COVER AND CONSTANCY. Survey area with 551 samples. Analysis limited to 394 samples with 25% cover. Tabulation includes species with constancy 25%. n = number of samples. Cov = % cover within the group. Con = % constancy. ' Pooled from species “‘misc’’ (species found in <5 samples), and ““PLspp”’ (Plantago spp.). 7 Pooled from Bromus diandrus, B. hordaceus, B. rubens, Lolium multiflorum, and HOspp (Hordeum spp.). * All cover, including species with <5% constancy. TRMA PUHO SCBO/misc DISP/glyco DISP/halo JUBU See n=33 n=19 n=19 n=61 n=43 £42.91 =44 Cov Cov Con Cov Con Cov Con Cov Con Cov Con Cov Con Triglochin maritima 1.0 14 100 1 7 Puccinellia howellii 10.8 1 1D 28 100 6 32 i “23 Low frequency glycophytes! 0.8 13 OS pee NS) Scribneria bolanderi 0.6 8 37 Annual grasses? es) 1 5 10 63 2 il Distichlis spicata 8.6 2 26 54 5 97 29 98 1 16 Polypogon monspeliensis 0.7 2 tit 6 4] <1 7 Juncus bufonius 1.4 1 21 D2 20 15 100 Total cover? De) 15 30 26 66 46 18 by 0.5, “‘saturated”’ by 0.75, and “‘standing water’”’ by 1.0. The resulting composite values ranged from 25 to 100. A Mann-Whitney test (Minitab 1991) was used to compare the moisture index values of the Puccinellia and Distichlis samples. Species monitoring. Following vegetation anal- ysis, groups of four 1 X 2 meter plots were as- signed to monitor the biotic and edaphic conditions of each of ten vegetation categories of interest, dur- ing the period from June 1993 to December 1994. To attempt to identify the conditions responsible for the variability of the Puccinellia stands, plots were assigned to monitor six stature and density combi- nations found within the population. The remaining sixteen plots were assigned to monitor nearly pure stands of the three next largest contributors to cov- er, Distichlis, Triglochin, Juncus bufonius, and also to barren locations. Initially the plots were random- ly assigned to suitable locations identified by the systematic sampling; however, this produced a poor spatial distribution within and between the springs, therefore some plots were reassigned to other ran- dom or nearby locations. Soil samples were analyzed for texture and rock fragment content, and for soluble and exchangeable cations, nitrogen, phosphorus, and organic matter. Soil moisture, salinity, pH, growth and cover were monitored approximately monthly during the peri- od. In the case of Puccinellia, additional biotic re- sponses were monitored, such as culm and seed production and seedling survival. This paper re- ports only salinity and cover. Except for salinity, and to a lesser extent, soil moisture, little difference was noted between the conditions associated with the monitored groups. (Among the Puccinellia groups, stature was inversely proportional to rock fragments.) The plots from the high density/medi- um stature ““H3”’’ group will be used to represent Puccinellia, because this group seemed to reflect conditions favorable for growth, and the vegetation in the four plots remained stable for most of the duration of the study. At each monitoring visit, several soil samples were collected from the periphery of each plot and pooled to form a composite weighing 300—600 g. After air-drying, 100 g portions of the fine-earth fraction were used to determine salinity by the sat- urated-extract procedure (Roades 1982). The ratio of the water content of the saturated paste to that of the original soil sample was used to convert the electroconductivity to a value presumed equivalent to the original field salinity. For specimens contain- ing precipitated salt at the time of collection, this method produced artificial values exceeding the conductivity of a saturated solution (e.g., 226 dS/ M at 25°C for NaCl), but nonetheless roughly pro- portional to the salt content. Greenhouse. In the greenhouse, Puccinellia growth was monitored at five levels of salinity, ranging from 0 to 80 dS/m, using 4 cm Puccinellia plants that had been germinated and reared at 20 dS/m, a typical field level. The treatment groups were adjusted to their target level at a rate of 10 dS/m per 8 days. Germination was also monitored under a similar range of salinities. RESULTS AND DISCUSSION Vegetation analysis, association. According to the point-intercept data, the total vegetative cover was 25.2%, with Puccinellia and Distichlis 10.8% and 8.6%, respectively. Table 1 lists the cover of the major contributors, and summarizes the mean cover and constancy for the component species of each of the six vegetation groups suggested by TWINSPAN. The dendrogram in Figure 2 shows the sequence of the divisions. TWINSPAN initially segregated two largely monotypic groups from the rest of the data, TRMA (Triglochin), and PUHO, the latter capturing 94% of the Puccinellia cover. 2002] Level O 1 227 PUHO/TRMA 2 3 33 194 4 19 TRMA PUHO SCBO/misc Fic. 2. LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING 181 394 167 123 104 DISP 44 61 43 DISP/glyco DISP/halo JUBU TWINSPAN species dendrogram for 394 vegetation samples. Associations: TRMA = Triglochin maritima; PUHO = Puccinellia howellii; SCBO/misc = Scribneria bolanderi, Plantago spp., and miscellaneous low frequency species; DISP/glyco = higher-density Distichlis spicata sometimes associated with annual grasses; DISP/halo = lower- density Distichlis sometimes associated with Puccinellia and Polypogon; JUBU = Juncus bufonius. The distinction between TRMA and PUHO, how- ever, was not visually apparent in the field, where the wetter habitat seemed to support both species intermixed. Therefore, it seemed reasonable to re- gard the two as a single association, PUHO/TRMA. Distichlis was divided between two groups. Ap- proximately two-thirds of the cover was placed in a group that we labeled DISP/glyco, characterized by higher density Distichlis, and often accompanied by annual grasses and other glycophytes that are active during the rainy season. Puccinellia was no- tably absent. During the vegetation survey, we re- garded the annual grasses as presumably salt-tol- erant contaminants, but we revised this view later as we observed Distichlis emerge amid the brown stalks of annual grasses in areas beyond the as- sumed boundary of the salt spring. In the down- slope portions of these peripheral and sometimes disjunct zones there were seepages that had not been visible. The accumulating mineral stains and evaporate indicated that during summer, after the cessation of precipitation, salt water was able to reach the surface. We inferred that in the central Spring area, the presence of glycophyte grasses amid the Distichlis might also indicate a salinity regime that alternates between fresh in winter and saline in summer. The remaining % of the Distichlis, labeled DISP/ halo, had a lower mean density and a smaller gly- cophytic component. Puccinellia and/or Polypogon monspeliensis (L.) Desf. were sometimes present. Although the latter is not usually regarded as a hal- ophyte, it will germinate simultaneously with Puc- cinellia when seed of both species are irrigated di- rectly by salt spring surface flow (Levine personal observation). The primarily monotypic JUBU, capturing most of the Juncus bufonius, also had a small Puccinellia component. The heterogeneous SCBO/misc., in- cluding nearly all the Scribneria bolanderi (Thur- ber) Hackel and most of the miscellaneous low- frequency glycophytes, consisted of the remaining samples. Vegetation analysis, moisture. The vegetation sampling was performed well after the cessation of seasonal precipitation, when the surface moisture primarily reflected the influence of the spring dis- charge, but before the evaporative effects of sum- mer heat. Among the 235 samples predominated by a single species, the wetter samples were common- ly dominated by Puccinellia or Triglochin, the drier by Juncus and Scribneria, and Distichlis was inter- mediate (Fig. 3). A Mann-Whitney test showed that the Puccinellia samples had significantly greater surface moisture values than the Distichlis (P < 0.0001). Species monitoring. Possible insights into the re- lationship between growth and the salinity regime may be provided by examining the data from the monitoring plots for the four species (Fig. 4a—h). Although the sample size is small, the data suggest that areas occupied by different species may also 182 MADRONO [Vol. 49 samples with one species predominant surface moisture histograms f=] SCBO J PUHO[__]DISP JUBU WY TRMA sample count 25 35 45 55 moisture index center mark Fic. 3. 65 15 85 95 Histogram of surface moisture for the 235 vegetation samples 215% cover, where one species contributed more than 50% (the large majority contributing >66%). The dominant species are indicated by the acronyms defined in Figure 2. Values range from 25 = “dry” to 100 = “standing water’’. differ in the annual pattern of salinity, and by in- ference, the amount of irrigation. In the Puccinellia and Triglochin plots (Fig. 4a, c), saline conditions occurred year-round, and rose only moderately during the summer. Apparently, the volume of irrigation was sufficient to minimize the seasonal influence of precipitation or evapora- tion. In contrast, salinity in the Juncus plots and most of the Distichlis plots (Fig. 4e, g) was low from fall to spring, and became high during the summer. The exception was Distichlis plot #3, which remained saline through the winter. This plot was selected for monitoring because of the atypi- cally short stature and low density, and the adja- cency of robust Puccinellia, perhaps suggestive of the TWINSPAN group, DISP/halo. The growth cycles of the four species appear to track the change in salinity. Puccinellia (Fig. 4b) has a phenology similar to the glycophytic grasses of the region. Growth occurs during the cool months, and seed is produced before a summer dor- mancy. However, regeneration by germination or resprouting often begins in advance of the fall rains. Triglochin (Fig. 4d) remains active through the summer, but declines during the cold months. Its succulent foliage is susceptible to damage from freezing. Distichlis (Fig. 4f) typically dies back completely during the cold months, but its prefer- ence for warm weather and its high salt tolerance (Ungar 1974; Hansen et al. 1976) enables it to grow on sites that during the summer become too saline for Puccinellia. Salt build-up in the Juncus bufon- ius plots (Fig. 4h) was more extreme than in the Distichlis plots. An early-flowering annual, Juncus can complete its life cycle within a short period of low salinity during winter and spring. Though the ability of Distichlis to tolerate high salinity is by no means unique, its often greater => Fic. 4. Seasonal salinity and cover in sets of monitoring plots. a, b) H3-Puccinellia = high density, medium stature Puccinellia howellii. Reported conductivities are the saturated extract values adjusted for field moisture. Samples with accumulated salt produced artificial conductivity values that exceeded the level of a saturated solution (e.g., 226 dS/m at 20°C for NaCl), but which remained proportional to the salt content. 50 dS/m is approximately equivalent to coastal sea water. The unconnected cover values labeled “‘green + brown,” and “‘brown”’ were measured at the end of the previous growing season. LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING Species Monitoring Plots - June 1993 to December 1994 2002] a_H8-Puccinellia Estimated Salinity 100 —= PLOT-1 ~*~ PLOT-2 ~*~ PLOT-3 --a- PLOT-4 80 60 £ ” me) 40 20 dS/m dS/m dS/m Ja4-94 Mr6é My1 JI3 Se4 Oc 29 fe) Jn29-93 Se5 Nod Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oct No22 C __Triglochin Estimated Salinity 100 80 60 40 0 Jn29-93 Se5 NoS5 Ja4-94 Mr6 My1 JI3 Se4 Oc29 Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oc1 No22 e Distichlis Estimated Salinity 400 D-1 121 D-3 103 80 60 40 Ja4-94 M Myt JIS Se4 Oc29 No22 ie) Jn29-93 Sed No5 Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oct Juncus Estimated Salinity +2 491 J-2 983 J-1 489 100 80 40 Se4 Oc29 ie) Jn29-93 Sed NoS Ja4-94 Mré6 My1 Ji3 Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oc1 No22 b H8-Puccinellia % Green Cover 100 80 x (green+ brown) 60 40 0 Je24-93 Se10 No15 Fe23-94 My3 Jy13 Se22 De21 Au4 Oc7 De31 Mr31 Jn7 Au15 No3 d Triglochin % Green Cover 100 80 60 40 eres Fe23-94 My3 Jy13 Se22 De21 ie) Je24-93 Se10 No15 Au4 Oc7 De31 Mr31 Jn7 Au15 No3 f Distichlis % Green Cove 100 — — 80 60 40 0 = " Pres = “- = - Je24-93. Se10 Noi5 Fe23-04 My3 Jy13 Se22.—s- De? Au4 Oc7 De31 Mr31 Jn7 Au15 No3 h Juncus % Green Cover 100 80 60 =< 40 (brown) Jy13 Se22 De21 Fe23-94 My3 Au4 Oc7 De31 Mr31 Jn7 Au15 No3 6 . Je24-93 Se10 No15 183 184 MADRONO capacity compared with other species has been not- ed in other settings. For example, in New England, Distichlis rapidly re-colonized bare patches of coastal salt marsh that were created by shading from wrack (Bertness et al. 1992; Shumway 1995). These areas then become hypersaline due to in- creased evaporation. Their studies suggest that Dis- tichlis can invade and occupy patches too saline for the establishment and persistence of discrete indi- viduals by transporting along its runners water and carbon from ramets outside the hypersaline zone. The distribution of Distichlis within the salt springs seems to reflect not only its relatively high salt tolerance, but also a limited capacity for im- mersion. On both coasts Distichlis is a component of high marsh vegetation, where it is not subject to daily flooding (Macdonald and Barbour 1974; Sil- berhorn 1982; Macdonald 1988). In inland settings it is associated with only moderate moisture (Ungar 1974). The detrimental effect of inundation on Dis- tichlis, and the associated abiotic factors, was dem- onstrated in a Louisiana tidal marsh undergoing vegetation decline associated with relative sea level rise. Webb et al. (1995) transplanted sod blocks of four species level with the marsh surface and raised 20 cm higher, equivalent to the level of a nearby healthy marsh. All species produced less biomass at the ambient level, but the greatest reduction was in those typically associated with high marsh hab- itat, a twenty-fold difference in the case of Distich- lis. The poor growth at the ambient level was as- sociated with negative substrate redox potential at both 2 cm and 15 cm depth. The elevated plantings had uniformly higher potentials, with oxidized con- ditions at 2 cm. The comparatively larger number of plots as- signed to monitor Puccinellia, and their division into stand categories, helped provide examples of the variation in growing conditions. The timing of Puccinellia germination appeared related to sum- mer salinity. Seed in locations directly inundated by surface flow germinated as early as mid-sum- mer, while areas with high summer salinity pro- duced their major germination pulse in November or later. Locations with visible salt accumulation, including those near the evaporative edge of sur- face flows, seemed more prone to seedling mortal- ity, and to the failure of tufts to regenerate in the fall. The two Puccinellia plots located in the trans- plant area provided an example of salinity-related mortality, and evidence that the pattern of seepage can shift over time. During the first summer of monitoring, these high-density/high-stature stands had a measured salinity similar to Distichlis plots, and visible salt accumulation. At the same time, the plots were being invaded by Distichlis. The follow- ing fall, the Puccinellia tufts did not resprout, and though recruitment was heavy, little germination was recorded within the plot itself. In effect, the Puccinellia had regenerated, but had shifted in po- [Vol. 49 sition, and was no longer centered within the plot boundaries. Toward the end of the monitoring pe- riod, plot H3-3 provided a similar example of non- regeneration due to shifting irrigation (Fig. 4b), however, the visible salt accumulation within the plot was not reflected in the soil samples collected from the periphery. Dense germination occurred outside and adjacent to the plot boundary, where surface flow was evident. Greenhouse. The salinity preferences of Pucci- nellia are suggested by its behavior in the green- house. The greatest growth was seen in the group that remained at 20 dS/m. Growth at 60 dS/m was stunted, and 80 dS/m proved lethal. In the field, summer levels of 60 dS/m and higher were com- monly found in the Distichlis and Juncus plots, and also in the plots of low-density/low-stature Pucci- nellia. The amount and rate of germination was in- versely proportional to salinity, consistent with field observations that germination is delayed in areas that that accumulated salt during the previous sum- mer. CONCLUSIONS The large majority of the Puccinellia cover was found in monotypic stands. The Distichlis stands were divided into two groups. The larger portion, two-thirds of the total cover, contained no Pucci- nellia, but often supported annual grasses during the cool season. The presence of Puccinellia in the remaining one-third and in the Juncus stands, might indicate locations where habitat conditions were in- termediate or in flux. The vegetation at the salt springs can be divided into two groups, based on the relative influence of spring discharge. A saline-winter/saline-summer di- vision, typified by Puccinellia and Triglochin, tends to be wet year-round. The habitat remains saline during the rainy season, excluding glycophytes dur- ing the period of active Puccinellia growth. Salinity rises only moderately during the summer, permit- ting late-summer to early-fall germination by Puc- cinellia. A fresh-winter/saline-summer division, typified by Distichlis and Juncus, is more likely to occur at the periphery or at elevated areas within the springs, where the influence of the discharge is less direct. During the summer, wicking and evap- oration promote hypersalinity, but winter precipi- tation dilutes the salt sufficiently to permit the growth of glycophytes. The areas occupied by these vegetation types may shift somewhat over time in response to changes in the pattern of seepage and surface flow. This study concludes that a major requirement for Puccinellia, apart from suitable salinity, is a volume of spring discharge sufficient to maintain salinity during both the wet and dry seasons. In this region of relatively high precipitation, few salt springs may meet this requirement, but qualifying sites merit attention because they may harbor un- 2002] documented populations or could become candidate sites for outplanting. The evidence that Puccinellia and Distichlis have different hydrologic preferences provides useful information for managing the only known population of Puccinellia in the presence of Distichlis. For instance, Distichlis might be exclud- ed by lowering the grade sufficiently so that surface flow occurs year-round. This approach is currently being explored by CalTrans within the area of their transplant project. ACKNOWLEDGMENTS We wish to extend our appreciation to Sharon Stacey of CalTrans District 2, Redding, and to Gretchen Ring and Jennifer Gibson of the Whiskeytown National Recreation- al Area, for their assistance in this study, and for their continued commitment to the conservation of Puccinellia howellii. LITERATURE CITED Bacca, M. 1995. Control strategies to inhibit saltgrass (Distichlis spicata) encroachment upon Howell’s al- kali grass (Puccinellia howellii) at an inland miner- alized spring area in Shasta County, California. M.S. thesis. Humboldt State University, Arcata, CA. BERTNESS, M. D., L. GOUGH, AND S. W. SHUMWAY. 1992. Salt tolerances and the distribution of fugitive marsh plants. Ecology 73:1842-—1851. CALIFORNIA NATIVE PLANT SOCIETY (CNPS). 2001. Inven- tory of rare and endangered plants of California, sixth edition. Rare Plant Scientific advisory Committee, D. Tibor, convening editor. California Native Plant So- ciety, Sacramento, CA. CH2M HILL, 1991-1992. Reports of inorganic analyses of water samples supplied by CalTrans, lab numbers 31306, 32495, 32650, 32818, 32948, 33641. Avail- able at CalTrans, Redding, CA. Davis, J. I. 1990. Puccinellia howellii (Poaceae), a new species from California. Madrono 37:55-—58. FULGHAM, K. O., L. LEVINE, AND M. Bacca. 1997. Aut- ecological Study of Puccinellia howellii: contract 02E326 final report. Available at CalTrans, Redding, CA. HANSEN, D. J., P- DAYANANDAN, P. B. KAUFMAN, AND J. D. LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING 185 BROTHERSON. 1976. Ecological adaptations of salt marsh grass, Distichlis spicata (Gramineae) and en- vironmental factors affecting its growth and distri- bution. American Journal of Botany 63:635—650. Hitt, M. O., C. T: FE TER BRAAK, O. E R. VAN TONGEREN, AND J. B. BirKs. 1988. TWINSPAN. Modification by J. B. Birks of a FORTRAN program originally writ- ten by M.O. Hill, 1979. Available from U.K. Mirror Service. September 22, 2002. http://www.mirror.ac. uk/collections/Mensa-micros/local/msdos/science/ cornell_ecology/cep2.zip KARTESZ, J. T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. /n J. T. Kartesz and C. A. Meacham (eds.), Synthesis of the North American Flora, Version 1.0. North Carolina Botan- ical Garden, Chapel Hill, NC. MACDONALD, K. B. 1988. Coastal salt marsh. Pp. 263—294 in M. G. Barbour and J. Major (eds.), Terrestrial veg- etation of California. California Native Plant Society, Sacramento, CA. AND M. G. BARBourR. 1974. Beach and salt marsh vegetation of the North American Pacific coast. Pp. 175-235 in R. J. Reimold and W. H. Queen (eds.), Ecology of halophytes. Academic Press, New York, NY. MIniTaBs. 1991. MINITAB version 8.2, statistical software. Minitab, Inc., State College, PA. RoaDEs, J. D. 1982. Soluble salts. Pp. 167—180 in A. L. Page (ed.), Methods of soil analysis, Part 2, Chemical and microbiological properties, 2nd ed. American So- ciety of Agronomy, Inc., and Soil Science Society of America, Inc., Madison, WI. SHUMWAY, S. W. 1995. Physiological integration among clonal ramets during invasion of disturbance in a New England salt marsh. Annals of Botany 76:225—233. SILBERHORN, G. M. 1982. Common plants of the Mid- Atlantic coast. Johns Hopkins University Press, Bal- timore, MD. UnGar, I. A. 1974. Inland halophytes of the United States. Pp. 235-306 in R. J. Reimold and W. H. Queen (eds.), Ecology of halophytes. Academic Press, New York, NY. WEBB, E. C., I. A. MENDELSSOHN, AND B. J. WILSEY. 1995. Causes of vegetation dieback in a Louisiana salt marsh: a bioassay approach. Aquatic Biology 51: 281-289. MADRONO, Vol. 49, No. 3, pp. 186-188, 2002 SPHAGNUM BALTICUM IN A SOUTHERN ROCKY MOUNTAIN IRON FEN DAVID J. COOPER Department of Earth Resources, Colorado State University, Fort Collins, CO 80523 dcooper @rm.incc.net RICHARD E. ANDRUS Environmental Studies Program, Binghamton University, Binghamton, NY 13902 CHRISTOPHER D. ARP Department of Biology, Utah State University, Logan, UT 84322 Sphagnum balticum (Russow) C. Jensen is a widespread Holarctic peat moss of raised and blan- ket bogs, occurring partly or completely submerged in pools, in floating mats and on hummocks. It is known from arctic and subarctic Greenland, North America (Crum and Andersen 1981; Crum 1984), Scandinavia, the British Isles, Russia and northern China. It is one of the rarest plants in the United Kingdom (UK Biodiversity Group 1998). Until now, the known range of S. balticum in North America extended south to about 57 degrees north latitude in western and central Canada and it is un- known in the lower 48 United States. Therefore, it is striking to discover this species in the San Juan Mountains in southwestern Colorado disjunct by more than 2000 km from the main range of the species in northern Canada and Alaska (Sphagnum balticum, Cooper #2281, COLO, BING). During a regional analysis of iron fens, we found Sphagnum balticum in the Chattanooga iron fen (2990 m elevation, Latitude 37°50'N, Longitude 107°43’W) south of Red Mountain Pass where it is the most abundant moss in shallow pools, growing with the sedges Carex aquatilis Wahlenberg and C. utriculata Boott. It is easily separated in the field from the only other Sphagnum in section Cuspidata present in the area, S. angustifolium (Russow) C. Jens. by a laxer habit and conspicuous lingulate and spreading stem leaves. The fen is almost complete- ly covered by Sphagnum mats and hummocks, with abundant S. angustifolium, S. russowii Warnst., S. fuscum (Schimp.) Klinggr. and S. fimbriatum Wils. (nomenclature follows Crum 1984). We found S. girgensohnit Russow, another new Colorado re- cord, in a different iron fen. The water and soil in most Sphagnum-dominated peatlands is acidic. The origin of the acids has been linked to Sphagnum cation exchange capacity (Cly- mo 1963; Clymo and Hayward 1982; Glaser 1987), atmospheric acid deposition (Gorham 1967), bio- logical uptake of nutrient cations by plants (Mitsch and Gosselink 1994), and the buildup of organic acids by decomposition (Gorham et al. 1984). These processes control autochthonous production and accumulation of acids in ombrogenous peat- lands (bogs). Fens in the Rocky Mountains, how- ever are soligenous (formed on slopes) or limno- genous (formed on lake margins), and are season- ally flushed with abundant snow melt water (Coo- per and Andrus 1994). Because strong flushing occurs, the pH of surface and ground water is con- trolled by the chemistry of watershed surface and ground waters, not autochthonous acid production. Warm and dry summers with long rainless peri- ods and a large evaporative demand are character- istic of the continental climate of the southern Rocky Mountains. Peatlands occur only where a continuous supply of ground water maintains pe- rennially saturated soil conditions. The most com- mon peatlands are transitional fens with slightly acid waters in mountain ranges of granite, rhyolite or metamorphic bedrock (Cooper and Andrus 1994) and extreme rich fens with circumneutral or basic waters in areas of, limestone, dolomite, shale, and basalt bedrock (Cooper 1996). Colorado iron fens appear to be geochemically unique and occur in areas with highly mineralized outcrops, such as occur on Red Mountain Pass. While iron fens are hydrologically similar to other Colorado fens, with ground water discharge per- manently saturating valley bottom wetlands, the source water flows through fractured iron pyrite- rich bedrock and talus, oxidizing pyrite and form- ing sulfuric acid. Chattanooga iron fen surface water has a pH from 3.8 to 4.4, similar to bogs and poor fens in northern Minnesota (Glaser 1987). Acid drainage from historical metal mines in Colorado produces similar low pH waters, and pollutes some Colorado wetlands (Arp et al. 1999), however iron fens are natural ecosystems. Although geochemical process- es external to the peatland generate the acids in iron fens, the fen flora is limited to acid tolerant species that also occur in poor fens and bogs. Bog waters have low ion concentrations because precipitation is the primary water supply. In con- trast, iron fens have high ion concentrations be- cause sulfuric acid produced in the watershed leaches and mobilizes metal and base cations from rock. Concentrations of Ca?* in Chattanooga fen 2002] surface water range from 14—20 mg/liter, and SO, averages 117 mg/liter. Ca** concentration in bogs range from 0.5 to 2.0 mg/liter (Glaser et al. 1981, 1990), in poor fens from 2.0—5.0 mg/liter (Sjérs 1963: Glaser et al. 1981), transitional fens in the Rocky Mountains from 2.0—7.0 mg/liter (Cooper and Andrus 1994), and rich fens from 5.0—30.0 mg/ liter (Glaser et al. 1981, 1990). Thus, the Chatta- nooga iron fen has Ca** concentration most similar to a rich fen. High iron concentrations (0.2—6.3 mg/ liter) precipitate onto fen organic matter forming terraced bog iron ore (limonite) deposits (Harrer and Tesch 1959), which are characteristic of iron fens. Limonite terraces perch the water table and form extensive networks of pools and ponds that are uncommon in most other Colorado fens. The presence of plants, such as Sphagnum bal- ticum, that are widely disjunct from their main ranges has always intrigued biogeographers. Did people, migratory birds or other vectors transport these plants? Are they the last vestiges of plant populations that established in the southern Rocky Mountains from the north during the Pleistocene as suggested by Hooker and Gray (1880)? Or do they reflect much older plant evolutionary and devel- opmental patterns as suggested by Weber (1965)? It is unlikely that animals, people or wind would have dispersed Sphagnum balticum, and Colorado has numerous other boreal montane bryophytes and lichens that exist in tiny populations disjunct from their main ranges in the boreal and subarctic re- gions of North America. Examples include Cladina stellaris (Opiz) Brodo found in a few 10 m? patches on the margin of a fen in the Tarryall Range, Sphagnum platyphyllum (Braithwaite) Warnstorf in two small wetlands in the Sawatch Range, S. con- tortum Schultz in one small wetland in the Front Range, Paludella squarrosa (Hedwig) Bridel in a couple of tiny alpine wetlands on Guanella Pass, and Scorpidium scorpioides (Hedwig) Limpricht in a few acres of calcareous fen in South Park (Cooper 1991; Weber and Wittmann 1996). The large num- ber of species persisting as localized populations, many found in only one fen, or on one mountain slope, suggests that these populations are the ves- tiges of what may have been wider distributions along the Cordillera that are much older than the Pleistocene. Many Colorado fens have basal '*C dates of 10,000—12,000 years BP (Cooper 1990; Chimner and Cooper 2002), with peat body initia- tion soon after the melting of Pleistocene glaciers. Boreal montane species could have found stable re- fugia in the southern Rocky Mountains and per- sisted through the Holocene. It is striking that new populations of widespread and well-known boreal montane plants are still be- ing found in Colorado, a state with modern floristic manuals (e.g., Weber and Wittmann 2001), numer- ous botanists, and a rich history of botanical inves- tigation. It indicates that additional species are like- ly to be discovered. Rare plants, such as S. balti- COOPER ET AL.: Sphagnum balticum in the Sourthern Rocky Mountains 187 cum, occur in very specialized habitats, such as iron fens, that are difficult to find. Careful inventories are necessary to make certain that logging, mining, and recreation and water developments do not in- advertently destroy the only populations of species occurring in the southern Rocky Mountains, and that may have existed here for many millennia. It also indicates the sensitivity of these habitats and populations to hydrologic variability that could ac- company human induced climate changes. Since populations of these species are very small and the circumstances of survival of each species are likely to be unique, there is no chance for species replen- ishment. ACKNOWLEDGMENTS This research was funded by a grant from the State of Colorado Natural Areas Program. We thank Janet Coles, Kathy Carsey, Gay Austin and Paula Lehr for their assis- tance in the field, and for Dr. W. A. Weber for discussions on biogeography. The identification of S. balticum was verified by Dr. Kjell Flatberg, University of Trondheim, Norway, and we thank him for his assistance. We also thank three reviewers for comments, which improved this manuscript. LITERATURE CITED Arp, C. D., D. J. COOPER, AND J. D. STEDNICK. 1999. The effects of acid rock drainage on Carex aquatilis leaf litter decomposition in Rocky Mountain fens. Wet- lands 19:665—674. CLymo, R. S. 1963. Ion exchange in Sphagnum and its relation to bog ecology. Annals of Botany 27:309-— 324. CLymo, R. S. AND P. M. HAywarpD. 1982. The ecology of Sphagnum. Pp. 229-289 in A. J. E. Smith (ed.), Bry- ophtye ecology. Chapman and Hall, London, U.K. CHIMNER, R. A. AND D. J. COOPER. 2002. Modeling carbon accumulation in fens using the century ecosystem model. Wetlands 22:100—110. Cooper, D. J. 1990. The ecology of wetlands in Big Meadows, Rocky Mountain National Park, Colorado: the correlation of vegetation, soils and hydrology. Bi- ological Report 90(15). U.S. Department of the In- terior, Fish and Wildlife Service, Washington, DC. . 1991. The habitats of three boreal fen mosses new to the southern Rocky Mountains of Colorado. The Bryologist 94:49—50. . 1996. Water and soil chemistry, floristics and phytosociology of the extreme rich High Creek fen, in South Park, Colorado, U.S.A. Canadian Journal of Botany 74:1801—1811. AND R. E. ANpDRuS. 1994. Patterns of vegetation and water chemistry in peatlands of the west-central Wind River Range, Wyoming, U.S.A. Canadian Jour- nal of Botany 72:1586—1597. Crum, H. 1984. North American flora: Sphagnopsida, Sphagnaceae. New York Botanical Garden, New York, NY. AND L. E. ANDERSEN. 1981. Mosses of eastern North America. Columbia University Press, New York, NY. GLASER, P. H. 1987. The ecology of patterned boreal peat- lands of northern Minnesota: a community profile. 188 MADRONO Biological Report 85(7.14). U.S. Department of In- terior, Fish and Wildlife Service, Washington, DC. , G. A. WHEELER, E. GORHAM, AND H. E. WRIGHT, JR. 1981. The patterned mires of the Red Lake Peat- land, northern Minnesota: vegetation, water chemis- try, and landforms. Journal of Ecology 69:575-—599. , J. A. JANSSENS, AND D. I. SIEGEL. 1990. The re- sponse of vegetation to chemical and hydrological gradients in the Lost River Peatland, northern Min- nesota. Journal of Ecology 78:1021—1048. GORHAM, E. 1967. Some chemical aspects of wetland ecology. Committee on Geotechnical Research, Na- tional Research Council on Canada, No. 90, pp. 20— 38. , 8. E. BAYLEY, AND D. W. SCHINDLER. 1984. Eco- logical effects of acid deposition on peatlands: a ne- glected field in “‘acid-rain” research. Canadian Jour- nal of Fisheries and Aquatic Science 41:1256—1268. HARRER, C. M. AND W. J. TESCH, JR. 1959. Reconnaissance of iron occurrences in Colorado. U.S. Department of [Vol. 49 the Interior, Bureau of Mines, Information Circular 7918. HOOKER, J. D. AND A. GRAY. 1880. The vegetation of the Rocky Mountain region and a comparison with that of other parts of the world. U.S. Geological Survey Territories 6:1—62. Mitscu, W. J. AND J. G. GOSSELINK. 1994. Wetlands, 2nd ed. Van Nostrand Reinhold, New York, NY. Syors, H. 1963. Bogs and fens on Attawapiskat River, northern Ontario. Bulletin of National Museum of Canada 186:45—103. UK Bropiversity Group. 1998. Tranche 2 action plans, plants and fungi. English Nature, London, U.K. WEBER, W. A. 1965. Plant geography of the southern Rocky Mountains. Pp. 453—468 in H. E. Wright, Jr. and David G. Frey (eds.), The Quaternary of the Unit- ed States. Princeton University Press, Princeton, NJ. AND R. C. WITTMANN. 2001. Colorado flora: east- ern slope, revised edition. University Press of Colo- rado, Niwot, CO. MADRONO, Vol. 49, No. 3, pp. 189-192, 2002 EVIDENCE OF A NOVEL LINEAGE WITHIN THE PONDEROSAE ANN M. PATTEN! AND STEVEN J. BRUNSFELD? Department of Forest Resources, University of Idaho, Moscow, ID USA 83843 ABSTRACT Phylogenetic analysis of the DNA of a putative portion of the nuclear NADH-specific nitrate reductase gene revealed the existence of a Pinus jeffreyi lineage that gave rise to P. washoensis and the North Plateau race of P. ponderosa var. ponderosa. These data are consistent with Lauria’s (1991) hypotheses that the North Plateau race is genetically distinct from the other races of the species, and that this race should be considered conspecific with P. washoensis. Pinus subsection Ponderosae is an economically important and well-represented group across much of western North America. However, the species in this group have been the source of considerable tax- onomic disagreement (Lauria 1991, 1997; Kral 1993; Rehfeldt 1999). Taxonomic treatments and inferences about the evolution of the Ponderosae have been based on a large number of different data sets, including quantitative morphological charac- ters (Peloquin 1984; Rehfeldt et al. 1996; Rehfeldt 1999), terpene chemistry (Mirov 1961; Smith 1964, 1967, 1977; von Rudloff and Lapp 1991), isozymes (Niebling and Conkle 1990), crossability (Critch- field 1984), provenance analysis (Wells 1964; van Haverbeke 1986), and the fossil record (Stockey 1984; Axelrod 1986). Some of these studies have produced conflicting data or additional uncertainty because of the omission of key species or varieties (Lauria 1991). The primary objective of this limited study was to test two of Lauria’s hypotheses: (1) the North Plateau race of ponderosa pine (Pinus ponderosa Douglas ex Lawson and C. Lawson) is a distinct genetic entity relative to the other races of this spe- cies (Lauria 1991); and (2) Washoe pine (Pinus washoensis Mason and Stockwell) and the North Plateau race of ponderosa pine are conspecific (Lauria 1997). Sampling thus focused primarily on the five geographic races of ponderosa pine (Smith 1977; Conkle and Critchfield 1988), Washoe pine, and Jeffrey pine (Pinus jeffreyi Grev. and Balf.). Three other members of Ponderosae, Pinus arizon- ica Engelm. and Martinez, Pinus durangensis Mar- tinez, and Pinus engelmannii Carr., were included for comparison of relative genetic divergence. Pi- nus coulteri D. Don from subsection Sabinianae was also included because of its high crossability with Jeffrey pine (Zobel 1951) and the close rela- tionship between subsections Sabinianae and Pon- derosae exhibited in a chloroplast DNA analysis ' Current address: The Institute of Biological Chemis- try, Washington State University, Pullman, WA 99164- 6340. * Corresponding author. E-mail: sbruns @uidaho.edu. (Kupkin et al. 1996). Pinus contorta Dougl. ex Loud. was included as an outgroup. Most taxa were represented by two samples; however four samples from the North Plateau race and one sample from each of the three recognized populations of Washoe pine were included (Table 1). This study was con- ducted concurrently with a larger ecological genet- ics study (Rehfeldt 1999) of Washoe pine, Jeffrey pine, and ponderosa pine. DNA was isolated from needle tissue according to Lodhi et al. (1994). PCR products were gener- ated using primers designed to target a region cod- ing for the two hinges that connect the internal heme domain to the amino and carboxy terminal domains of the nuclear NADH-specific nitrate re- ductase (NADH-NR) gene (Zhou et al. 1995; Patten 1999). Manual sequencing of the PCR product was conducted using the USB Sequenase Kit (Amer- sham). A BLAST search of the GenBank data base did not reveal a match between our sequences and those reported as NADH-NR. However, the se- quence and structure of nitrate reductase is not known for any gymnosperm. Furthermore, the PCR primers are targeted to an area known to contain introns (Zhou and Kleinhofs 1996), lessening the chance of similarity to pine. Thus, to be conser- vative, the sequences might best be considered anonymous, although the parsimonious distribution of synapomorphies strongly suggests the sequences are orthologous (Fig. 1). Paralogous PCR products would likely exhibit a more random distribution with respect to taxonomic classification and geog- raphy (see below). Because there were variable amounts of missing data at the termini of the PCR products, a 287-bp fragment was used in the final phylogenetic anal- ysis. The sequences were analyzed using PAUP version 3.1.1 (Swofford 1993). The branch and bound exact algorithm was used and two most par- simonious trees were recovered. These differed only in the resolution of one dichotomy. Both trees had a length of 25 and a consistency index of 0.90. A consensus tree was constructed and all branches of zero length were collapsed (Fig. 1). This phylogenetic analysis offers novel insights 190 MADRONO [Vol. 49 TABLE 1. 27 SAMPLES USED IN PHYLOGENETIC ANALYSIS OF NORTH AMERICAN PONDEROSAE. Sample Location Collection GenBank # P. contorta Latah Co., ID A. Patten #95-14 AF06764 P. coulteri/A Black Mtn., CA A. Patten #95-10 U77801 P. coulteri/B Black Mtn., CA A. Patten #95-9 U77802 P. jeffreyi/A Nevada Co., CA USFS U77803 P. jeffreyi/B Nevada Co., CA USFS U77804 P. ponderosa/NP-ID Idaho Co., ID A. Patten #95-4 U77805 P. ponderosa/NP-OR Benton Co., OR G. Rehfeldt #94-9 U77810 P. ponderosa/NP-WA1 Kittitas Co., WA G. Rehfeldt #94-7 U77806 P. ponderosal[NP-WA2 Puget Sound, WA G. Rehfeldt #94-8 U77819 P. washoensis/BP Babbitt Peak, CA USES #55 U77807 P. washoensis/MR Mount Rose, NV USES #27 U77808 P. washoensis/[WM Warner Mtns., CA USES #48 U77809 P. ponderosa/RM-A Daggett Co., UT USES #206 U77 sil P. ponderosa/RM-B Fergus Co., MT C. Baldwin #95-12 U77820 P..ponderosa/PAC-A Washoe Co., NV USFS #60 U77814 P. ponderosa/PAC-B Nevada Co., CA USES #71 U77815 P. ponderosa/PAC-C Illinois River, OR C. Baldwin #95-2 U77818 P. ponderosa/SCA-A Black Mtn., CA A. Patten #95-8 U77816 P. ponderosa/SCA-B Fraser Park, CA C. Baldwin #95-13 U77817 P. ponderosalSW-A Lincoln Co., NM USFS #356 U77813 P. ponderosa/SW-B Graham Co., AZ USES #10-300 U77812 P. arizonica/US-A Graham Co., Az USES #1-9 WT7S22 P. arizonica/US-B Cochise Co., AZ USES #4-119 U77821 P. arizonica/MX Chihuahua, MX USFS #25-100 U77823 P. durangensis Durango, MX USFS #29-400 U77824 P. engelmannii/A Cochise Co., AZ USES #5-108 UZ7825 P. engelmannii/B Chihuahua, MX USFS #23-200 U77826 as well as support for previously proposed relation- ships within the Ponderosae. The most intriguing finding is the existence of a P. jeffreyi lineage, which contains P. jeffreyi in a basal position and P. washoensis and the North Plateau race of P. pon- derosa as derived taxa (Fig. 1). Based on this anal- P. contorta P. coulteri/A P. coulteri/B P. jettreyi/ A P. ponderosa /NP - ID P. washoensis /BP - CA P. ponderosa /NP - WA1 P. washoensis /MR - NV P. washoensis /WM - CA P. ponderosa /NP - OR P. jettreyi/B P. arizonica/US - B * * All other samples have zero branch length Fic. 1. Strict consensus tree for 27 samples of Pinus using branch and bound search and bootstrap of 1000 rep- licates in PAUP 3.1.1. Numbers above the branches rep- resent unambigious base substitutions, bracketed numbers represent indels. Numbers below the branches represent bootstrap values and decay indices, respectively. ysis, P. washoensis and the North Plateau race sam- ples share a common ancestor with sample A of P. Jeffreyi, and this monophyletic group is separated from sample B of P. jeffreyi by four derived nucle- otide substitutions. The marked differentiation be- tween the P. jeffreyi samples was unexpected be- cause they come from the same seed provenance in the Sierra Nevada. However, extant P. jeffreyi has been shown to possess rich intrapopulational ge- netic diversity (Furnier and Adams 1986). Future phylogenetic studies involving P. jeffreyi need to have considerably greater intra- and interpopula- tional sampling. Although the P. jeffreyi lineage de- scribed in this paper is novel, previous literature supports a close relationship among Jeffrey pine, Washoe pine, and the North Plateau race. Lauria (1991) observed that the purple color of immature ovulate cones of these three taxa is unique among the Ponderosae. All other members of this subsec- tion, including the geographically-proximal Pacific race of ponderosa pine, exhibit green to greenish- yellow ovulate cone color (Critchfield 1984). Mirov (1967) noted the similarity of ovulate cone structure between Washoe and Jeffrey pines. The results of the phylogenetic analysis also sug- gest that Washoe pine and the North Plateau race of ponderosa pine form a robust clade (bootstrap 100%), within which both taxa are polyphyletic (Fig. 1). The putative NADH-NR sequences from these samples appear to represent four alleles dis- tributed randomly among six populations. Due to 2002] the small sample size, it cannot be determined how these alleles are structured within or among popu- lations or species. Expanded studies using addition- al informative DNA regions and statistically-signif- icant intrapopulation sampling are needed. Regard- less, the random distribution of alleles is consistent with the close relationship or even conspecific sta- tus previously proposed between Washoe pine and the North Plateau race (Wells 1964; Haller 1965; Critchfield 1984; Niebling and Conkle 1990; Lauria 1991, 1997; Brayshaw 1996; Rehfeldt 1999). Prov- enance tests by Wells (1964) showed that Washoe pine was more similiar to the North Plateau race than it was to the Pacific race of ponderosa pine. Critchfield (1984) proposed that Washoe pine could be a recent derivative of the North Plateau race. This view is consistent with a close relationship inferred from isozyme data (Niebling and Conkle 1990). Washoe pine and the North Plateau race were found to have a genetic distance of 0.004, a value nearly identical to the genetic distance found among the three recognized populations of Washoe pine. Furthermore, Rehfeldt’s (1999) quantitative analysis of adaptive traits determined that Washoe pine and the North Plateau race of ponderosa pine are very closely related. Based on his and previous research, Rehfeldt (1999) concluded that these taxa should be considered synonymous. The high level of divergence of the Washoe/ North Plateau clade (seven synapomorphies and two deletions) suggests an origin involving small population size and isolation. The divergent Wash- oe pine/North Plateau clade eventually came to oc- cupy the Willamette Valley and the region approx- imating the current extent of the maritime climate east of the Cascade crest. Members of this clade dispersed as far east as the Continental Divide, where an abrupt genetic transition is evident. Latta and Mitton (1999) found a steep east-west cline in cpDNA and mtDNA, consistent with secondary contact between diverged taxa. Similarly, Critch- field (1984) previously proposed that Washoe pine expanded over the Pacific Northwest only to be lat- er absorbed by the North Plateau race of ponderosa pine. This study includes a single tree from a popula- tion on the Fort Lewis plains of the Puget Sound. This population is isolated from the North Plateau race by the Cascade mountains. Our DNA sequence from this individual is identical to that of Ponde- rosae found in the Rocky Mountains, Sierra Madre, and Sierra Nevada, suggesting that ponderosa pine from the Puget Sound region could be a relictual population of formerly widespread P. ponderosa s.l. This is in no way conclusive, as a single DNA marker from a single specimen may not be repre- sentative of a population. Nevertheless, it does sug- gest that the genetics of Ponderosae in the Puget Sound area needs to be investigated in detail as numerous studies have hypothesized that the Puget Sound region was a glacial (Pleistocene) refugium PATTEN AND BRUNSFELD: JEFFREY PINE LINEAGE ON for numerous plants and animals (e.g., Harris 1965; Steinhoff et al. 1983; Heusser 1985; Soltis et al. 1997). This study does not support the traditional vari- etal classification of ponderosa pine (e.g., Conkle and Critchfield 1988). Variety scopulorum, the Rocky Mountain form, is not distinct in our anal- ysis from the Pacific and the Southern California races of var. ponderosa. The principal finding of this phylogenetic analysis is the existence of a P. Jeffreyi lineage that gave rise to Washoe pine and the North Plateau race of var. ponderosa, a race that appears to be fundamentally distinct from the re- mainder of P. ponderosa s.l. Our results cannot confirm conspecificity of Washoe pine and the North Plateau race of ponderosa pine, but do indi- cate a very close relationship of these taxa to each other and to Jeffrey pine. This study did not lend insights into the relationships among the south- western and Mexican species of subsection Pon- derosae, suggesting relatively little genetic diver- gence of these taxa compared to the P. jeffreyi lin- eage. However, morphological and ecological data (e.g., Peloquin 1984; Rehfeldt et al. 1996) indicate the existence of significant patterns of genetic di- vergence in the Ponderosae of the southwestern U.S. and Mexico. We hope that this note provides the impetus for a more detailed phylogenetic study of the subsection Ponderosae. Larger sample sizes and multiple, rapidly-evolving DNA segments should be analyzed to test the results reported here. ACKNOWLEDGMENTS We thank G.E. Rehfeldt for providing samples and in- valuable guidance, Pam Soltis for assistance with the phy- logenetic analysis and her generous advice during the pro- ject, and Calib Baldwin for collecting several samples. This research was supported by funds provided by the Intermountain Research Station, Forest Service, U‘S. Dept. of Agriculture, a grant from the Stillinger Trust Fund, University of Idaho, and through funding received from the NSF-Idaho EPSCoR program under NSF Co- operative Agreement #OSR-9350539. 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Monoterpenes of ponderosa pine xylem resin in western United States. Technical Bulletin No. 1532, U.S.D.A. Forest Service, Pacific Southwest Forest and Range Experiment Station, Berkeley, CA. SoLTis, D. E., M. A. GITZENDANNER, D. D. STRENGE, AND P. S. Sortis. 1997. Chloroplast DNA intraspecific phylogeography of plants from the Pacific Northwest of North America. Plant and Systematic Evolution 206:353-—373. STEINHOFF, R. J., D. G. JOYCE, AND L. Fins. 1983. Isozyme variation in Pinus monticola. Canadian Journal of Forestry Research 13:1122—1132. STOCKEY, R. A. 1984. Middle Eocene Pinus remains from British Columbia. Botanical Gazette 145:262—274. SWOFFORD, D. L. 1993. PAUP: Phylogenetic analysis us- ing parsimony. Illinois Natural History Survey, Champaign, IL. VAN HAVERBEKE, D. FE 1986. Genetic variation in ponde- rosa pine: a 15-year test of provenances in the great plains. Rep. No. RM-265. Rocky Mountain Forest and Range Experiment Station U.S.D.A. Forest Ser- vice, Fort Collins, CO. VON RUDLOFF, E. AND M. S. LApp. 1991. Chemosystematic studies in the genus Pinus. VII. The leaf oil terpene composition of ponderosa pine, Pinus ponderosa. Ca- nadian Journal of Botany 70:374-—378. WELLS, O. O. 1964. Geographic variation in ponderosa pine. Silvae Genetica 13:89—103. ZHOU, J., A. KILIAN, R. WARNER, AND A. KLEINHOFS. 1995. Variation of nitrate reductase genes in selected grass species. Genome 38:919—927. AND A. KLEINHOFS. 1996. Molecular evolution of nitrate reductases. Journal of Molecular Evolution 42: 432-442. ZOBEL, B. 1951. The natural hybrid between Coulter and Jeffrey pines. Evolution 5:405—413. MADRONO, Vol. 49, No. 3, p. 193-197, 2002 NOTEWORTHY COLLECTIONS ALASKA ALLIARIA PETIOLATA (Bieb.) Cavara & Grande (BRAS- SICACEAE).—City and Borough of Juneau, Alaska, in the landscaped area next to a parking lot near the inter- section of Village and Wittier streets, 58°18'05’N, 134°24'53"W, 6 June 2001. One mature, flowering plant was found by P. Johnson and removed before it set seed. It is unclear how this plant arrived at this urban area. It is unlikely that it was introduced by the landscaping ac- tivities since there had been no recent additions of plants or soil. Previous knowledge. Native to northern Europe. Com- monly called garlic mustard because of the characteristic smell of its leaves when crushed, it is a highly competitive, aggressive herbaceous invader that forms dense understory populations. Present in 34 USA states (http:// plants.usda.gov), particularly in the eastern USA (Rollins, R.C., The Cruciferae of Continental North America. 1993. Stanford University Press) and the four Canadian provinces of British Columbia, New Brunswick, Ontario, and Quebec (http:/infoweb.magi.com/~ehaber/ipcan.html). The previ- ously known British Columbia collections were near Van- couver and Vernon (E. Haber personal communication). Significance. First record in Alaska. The previously known sites in British Columbia are approximately 1300 km to the southeast. Alliaria petiolata is one of the most pestiferous non-native invasive species of forest understo- ries in the USA and Canada. It can form monospecific stands, exclude native communities, and be essentially im- possible to eradicate once it is established. While it was originally believed this plant was the only individual growing in Alaska, P. Johnson subsequently found a large population of plants nearby. Although the early detection of this plant has been important, it is unclear whether the population can be successfully eradicated. The specimen has been placed in the Herbarium at the University of California, Davis (DAV). —Barry A. Rice, The Nature Conservancy, Wildland Invasive Species Team, Department of Vegetable Crops and Weed Sciences, University of California, Davis, CA 95616. —PHILLIP JOHNSON, P.O. Box 22898, Juneau, AK 99802. CALIFORNIA DROSERA ALICIAE Hamet (DROSERACEAE).—Mendo- cino county, CA, 39°15’N, 123°45’W, elevation 160 m, 2 November 1997. A few hundred meters west of Albion Little River Road, just south of the County Airport. A single spreading colony of plants was found in wet de- pressions and Sphagnum mounds in a pine/cypress pygmy forest. Previous knowledge. Native to South Africa, Drosera aliciae is a plant commonly grown by carnivorous plant enthusiasts. Significance. It has not previously been collected in California, and is probably a new introduction for North America. This plant was introduced by horticulturists with a number of other carnivorous plant taxa. It was repro- ducing both vegetatively and by seed. A more complete discussion is given under the Drosera capensis collection description. A specimen has been placed in the Herbarium at the University of California, Davis, #MR971101. DROSERA CAPENSIS L. (DROSERACEAE).—Mendocino county, CA, 39°15’N, 123°45’W, elevation 160 m, 2 No- vember 1997. A few hundred meters west of Albion Little River Road, just south of the County Airport. Large col- onies of plants were found growing in wet depressions and Sphagnum mounds in a pine/cypress pygmy forest. Previous knowledge. Native to South Africa, Drosera capensis is a common greenhouse weed in collections where carnivorous plants are grown. It is not listed in the various floristic works of California. The Jepson Manual notes the presence of D. linearis Goldie in Mendocino County, but this is probably an erroneous reference to ob- servations of D. capensis. A North American Drosera species, D. linearis does resemble D. capensis, but D. li- nearis is extremely difficult to cultivate and is nearly ab- sent from most collections. As such it is unlikely any hor- ticulturists have ever planted out D. linearis in California. D. linearis grows in Canada from Labrador west to On- tario, and in the US in Minnesota, Wisconsin, Michigan, and Maine. No plants of D. linearis were found in the area. Significance. This cluster of wildland locations has been used by carnivorous plant horticulturists for introduction experiments since the 1970s (P. D’Amato, Carnivorous Plant Newsletter, 1988, 17: 15-21). Plants from nearly every Carnivorous genus have been planted over the years, but most died within a few years. Non-native species that persisted and were spreading by seed or vegetative means were Drosera aliciae, D. binata, D. capensis, D. capillar- is, D. filiformis, D. intermedia, D. nitidula X occidentalis, Sarracenia flava, S. leucophylla, S. minor, S. purpurea, S. rubra, many interspecific Sarracenia hybrids, and Urtri- cularia subulata. Additional species present but which may be waifs were Dionaea muscipula, Drosera burman- nil, D. slackii, Pinguicula lusitanica, and Utricularia gib- ba. None of these plants are included in Californian flo- ristic works except for Darlingtonia californica, Drosera capensis (incorrectly listed in The Jepson Manual as D. linearis), D. filiformis, Sarracenia purpurea, and Utricu- laria gibba. Darlingtonia californica was abundant al- though it is not native to this location—its nearest natural occurrence is in central Trinity County (J. H. Rondeau, Carnivorous Plants of California, 1991, unpublished man- uscript). Many of the clumps of this plant were heavily damaged by infestations of greenhouse thrips, a condition not seen in natural populations. The only thriving species were Darlingtonia californica, Drosera capensis, and Utricularia subulata. Although it is unlikely any of these plants will spread from these isolated plantings, Drosera capensis or Utricularia subulata (which reproduce both vegetatively and by copious seed production) would be difficult to eradicate if they invaded other high quality natural habitats. The only carnivorous plant native to the site is Drosera rotundifolia. Those familiar with the site believe the Drosera rotundifolia is being displaced by the exotic species (C. Gardner personal communication). 194 Specimens have been placed in the Herbarium at the Uni- versity of California, Davis, #MR971103. UTRICULARIA SUBULATA L. (LENTIBULARIACEAE).— Mendocino county, CA, 39°15'N, 123°45’W, elevation 160 m, 2 November 1997. A few hundred meters west of Albion Little River Road, just south of the County Air- port. Large colonies of plants were found in wet depres- sions and water drainages in a pine/cypress pygmy forest. Previous knowledge. This is a widespread species found on every continent except Antarctica. In the United States it is found on the Atlantic and Gulf coasts from Massa- chusetts to Florida to Texas, and inland to Arkansas and Tennessee (P. Taylor, The Genus Utricularia: a Taxonomic Monograph, 1989, Kew Bulletin Additional Series XIV). It is a common greenhouse weed in collections of carniv- orous plants. Significance. A first collection for California. The plants were growing in densely matted clumps and were repro- ducing vegetatively and by seed. Both cleistogamous and chasmagomous flowers were present. Unless deliberately spread, it is unlikely this plant will escape from these plantings, but if it did it would be difficult to eradicate. Utricularia gibba (probably introduced) was also present in flower. A more complete discussion is given under the Drosera capensis collection description. A specimen of U. subulata has been placed in the Herbarium at the Uni- versity of California, Davis, #MR971102. —Barry A. RICE, The Nature Conservancy, Wildland Weeds Management and Research, Department of Vege- table Crops and Weed Sciences, University of California, Davis, CA 95616. OREGON AGROSTIS HOWELLIT Scribn. (POACEAE).—Linn Co., rare, N-sloping bench in a moist Acer circinatum-Carex deweyana community, Coburg Hills, 25 km NE of Eu- gene, T15S R2W S39, elev. 450 m, 3 Oct 1995, Brainerd 42 (OSC) (! K. L. Chambers 1996 OSC). Previous knowledge. Previously known as a narrow en- demic from a few sites on the south side of the Columbia River Gorge in northern Oregon. Significance. Circa 165 km SSW of previously docu- mented populations. CAREX SCIRPOIDEA Michx. subsp. STENOCHLAENA (Holm) A. Love & D. Léve (CYPERACEAE).—Lane Co., drip- ping cliff with Salix sitchensis, Agrostis, above Forest Ser- vice Road 19, near Cougar Dam, T16S RSE S31, elev. 520 m, 15 Jul 1998, Newhouse 98027 (MICH, OSC, WTU) (! A. A. Reznicek 2001 MICH). Previous knowledge. Ledge sedge ranges from Alaska south to Washington and Montana. Significance. First record for this subspecies in Oregon. DAPHNE LAUREOLA L. (THYMELAEACEAE).—Lane Co., Laurelwood golf course, Eugene, elev. 215 m, 13 Feb 1998, Newhouse 98002 (OSC); Hawkins Heights, Eugene, elev. 215 m, 20 Feb 1998, Newhouse 98003 (OSC); Ma- sonic Cemetery, Eugene, elev. 150 m, Aug 1997, New- house 97051 (OSC). Previous knowledge. Spurge-laurel is native to Europe, MADRONO [Vol. 49 and adventive in British Columbia and western Washing- ton, where it is bird-disseminated. Significance. First report as an escape from cultivation in Oregon. GALIUM PEDMONTANUM (Bellardi) All. (RUBI- ACEAE).—Benton Co., common in disturbed meadow 0.5 km NW of Pigeon Butte, elev. 80.m, T13S R5W S32, 5 Jun 1993, Zika 12025 (OSC, WTU). Previous knowledge. Mountain crosswort is native to the Mediterranean, and adventive in Idaho, Montana, and in the southeastern United States. Significance. First Oregon report; discovered by Robert Frenkel in 1992. PETASITES FRAGRANS (Vill.) C. Pres] (ASTERACEAE).— Benton Co., steep forested bank on W shore of Willamette River, Corvallis, elev. 60 m, T12S R5W S2, 3 Feb 1999, Zika 13717 (OSC, US, WTU); same site, 15 Mar 2000, Zika 14848 (OSC). Previous knowledge. Winter heliotrope is native to N Africa, and occasionally planted as an ornamental in west- ern Oregon. Significance. First report of an escape from cultivation in Oregon. —BRUCE NEWHOUSE and RICHARD BRAINERD, Salix As- sociates, 2525 Potter, Eugene, OR 97405; and PETER E ZIKA, Herbarium, Dept. of Botany, Box 355325, Univ. of Washington, Seattle, WA 98195-5325. OREGON ACAENA NOVAE-ZELANDIAE Kirk (ROSACEAE).—Coos Co., Randolph Road, near Route 101, 6 km N of Bandon, common weed on sandy banks, roadbeds, dikes, and cran- berry fields, with Crepis capillaris, Hypericum boreale, Juncus planifolius, J. canadensis, Poa annua, elev. 52 m, T28S R14W S4, 7 Sep 1999, Zika 14247 (OSC, WTU); Curry Co., Gold Beach, 13 Aug 1951, Jenkins s.n. (OSC); adventive on Azalea Lane, Wedderburn, elev. 30 m, T36S R1IS5W S25, 7 Jun 2000, Stansell 3196 (OSC); lawn weed, Route 101, Gold Beach Ranger Station, Gold Beach, T37S R1I5W S1, 17 Aug 2000, Stansell 3201 (OSC). Previous knowledge. Biddy-biddy is native to New Zea- land, and occasionally cultivated as a ground cover. It readily spreads via barbed fruits. Acaena is classified as a noxious weed in California, where it is found on disturbed ground along the coast. In Oregon it has been observed at several sites in addition to the ones vouchered, includ- ing a large population at Cape Blanco lighthouse in Curry Co. Significance. First report for Oregon. We first observed Acaena in June 1992 at the U.S. Forest Service office in Gold Beach, where lawn mowers scattered the seeds and led to an increase in the population. The Oregon Dept. of Agriculture has made several unsuccessful attempts to ex- tirpate the species with herbicides, starting in 1997. —Davip Pivorunas, Navy Region Southwest Natural Resource Office, 33000 Nixie Way, Bldg. 50, Suite 333, San Diego, CA 92147-5110; VEVA STANSELL, P.O. Box 6077, Pistol River, OR 97444-1575; and PETER FE ZIKA, Herbarium, Dept. of Botany, Box 355325, Univ. of Wash- ington, Seattle, WA 98195-5325. 2002] OREGON CERASTIUM PUMILUM Curtis (CARYOPHYLLA- CEAE).—Jackson Co., pasture with vernal pools, Route 234 E of Sams Valley, 12 May 1974, Chambers 3974 (OSC, WTU); Multnomah Co., silty shore, delta of Sandy River, elev. 4 m, 16 Apr 1992, Zika 11470 (OSC, WTU), Zika 11480 (WTU). Previous knowledge. Dwarf mouse-ear is native to Eu- rope, and naturalized in eastern North America as well as British Columbia. In the Pacific Northwest often growing among and confused with C. semidecandrum L. Significance. First report for Oregon. COTONEASTER DIVARICATUS Rehder & E.H. Wilson (RO- SACEAE).—Lane Co., bird-sown in thicket with Quercus garryana, Toxicodendron, Morse Ranch Park, Eugene, 30 Apr 1998, Love 9816 (OSC). Previous knowledge. Spreading cotoneaster is native to central China, and cultivated as an ornamental in the Pa- cific Northwest. Significance. First collection of an escape from culti- vation in Oregon. COTONEASTER INDURATUS J. Fryer & B. Hylm6 (ROSA- CEAE).—Lane Co., thickets, near Willow Creek, West Eugene, elev. 122 m, 28 May 1992, Zika 11593 (WTU); same population, 8 Jul 1997, Zika 13231 (WTU). Significance. First reports for hard cotoneaster as an es- cape from cultivation. Specimens identified by Jeanette Fryer. SORBUS CALIFORNICA Greene (CAPRIFOLIACEAE).— Klamath Co., Rim above Crater Lake, 12 Aug 1919, Sweetser s.n. (ORE): same site, elev. 2150 m, 13 Jul 1929, Wynd 1533 (ORE); Wizard Island, Crater Lake, 28 Jun 1934, Applegate 8977 (OSC); Phantom Ship, Crater Lake, elev. 1885 m, Zika 12516 (OSC). Significance. First collections for Oregon. All sites are in Crater Lake National Park. VERONICA VERNA L. (SCROPHULARIACEAE).—Union Co., sunny opening, Route 82 beside Wallowa River, 6 km SE of Minam, elev. 825 m, 2 Jun 1961, Mason 1142 (ORE): dirt road, Miller Flat, W shore of Wallow River, elev. 750 m, 21 May 1994, Zika 12182 (OSC); Wallowa Co., dirt road 11 km W of Enterprise, elev. 1220 m, 24 Apr 1961, Mason 807AA (OSC): 7 miles basalt cliff, Wal- lowa Falls, elev. 1525 m, 24 Jun 1962, Mason 5051 (OSC): Buck Creek near Imnaha River, elev. 580 m, 7 May 1991, Zika 11089 (OSC). Previous knowledge. Spring speedwell is native to Eu- rope. Crins et al. (Michigan Botanist 26: 161-166, 1987) discuss how to separate it from V. arvensis. Mason (1980, Guide to the Plants of the Wallowa Mountains of North- eastern Oregon, Museum of Natural History, Univ. of Oregon, Eugene) reported her collections of V. verna as V. triphyllos. The latter has lower leaves palmately lobed, and bracts shorter than fruiting pedicels. Veronica verna has pinnately lobed lower leaves and bracts longer than the fruiting pedicels. Significance. First documentation in Oregon. W ASHINGTON CARDAMINE FLEXUOSA With. (BRASSICACEAE).— Grays Harbor Co., moist ground, edge of building, near NOTEWORTHY COLLECTIONS 195 mouth of Boone Creek, elev. 5 m, 13 Dec 2001, Zika 16733 (WTU); King Co., wet sunny ditch, Burke Gilman Trail 2.4 km N of Matthews Beach, Lake City, Seattle, elev. 10 m, 30 Aug 2001, Zika 16467 & Jacobson (WTU);: weed in garden bed, Madison Park, Seattle, elev. 10 m, 31 Aug 2001, Zika 16474 (WTU); San Juan Co., wet ground in shade of Alnus rubra, Mineral Point, San Juan Island, elev. 5 m, 27 Oct 2001, Zika 16708A (WTU). Significance. First report for Washington for this Eur- asian native. CERASTIUM PUMILUM Curtis (CARYOPHYLLA- CEAE).—Island Co., dunes near W shore of Cranberry Lake, Whidby Island, elev. 3 m, 20 May 2000, Zika I5000A (WTU):; King Co., cracks in asphalt sidewalk, Montlake, Seattle, elev. 15 m, 20 May 2000, Zika 14998 (WTU);: San Juan Co., Turn Point, San Juan Island, 4 Apr 1992, Atkinson 307 (WTU): sand, Spencer Spit, Lopez Island, elev. 2 m, 21 May 2000, Zika 15005 (WTU). Significance. First report for Washington. COTONEASTER DIVARICATUS Rehder & E.H. Wilson (RO- SACEAE).—King Co., bird-sown in thickets, with Cory- lus cornuta, Gaultheria shallon, arboretum, near Union Bay, Seattle, elev. 20 m, 15 Sep 1999, Zika 14332 & Jacobson (WTU); same population, 26 Oct 2000, Zika 15609 (WTU; dupl. det. by Bertil Hylm6). Significance. First collection of an escape from culti- vation. COTONEASTER LUCIDUS Schltdl. (ROSACEAE).—Colum- bia Co., spreading and naturalized in Tucannon River bot- tomland, TON R41E S30, elev. 1045 m, 27 Jun 1989, Urban 89-001 (OSC). Previous knowledge. Shiny cotoneaster is native to Si- beria and Mongolia. It is occasionally cultivated in the Pacific Northwest, often under the misapplied name C. acutifolius Turcz. Significance. First collection of an escape from culti- vation. COTONEASTER NITENS Rehder & E.H. Wilson (ROSA- CEAE).—King Co., thickets, partial shade, campus of Univ. of Washington, Seattle, elev. 25 m, 26 Oct 1999, Zika 14660 & Jacobson (WTU); same population, 6 Nov 2000, Zika 15645 (WTU). Previous knowledge. Few-flowered cotoneaster is native to western China, and an uncommon ornamental in west- ern Washington. Significance. First collection of an escape from culti- vation. COTONEASTER SALICIFOLIUS Franch. (ROSACEAE).— King Co., spreading from cultivation to thickets, Kubota Gardens, Rainier Beach, Seattle, elev. 50 m, 3 Nov 1999, Zika 14704 (WTU); bird-sown in thickets, campus of Univ. of Washington, Seattle, elev. 25 m, 7 Nov 1999, Zika 14708 (WTU): bird-sown, partial shade, Madrona, Seattle, elev. 50 m, 2 Aug 2000, Zika 15187 (WTU): cracks in concrete wall, ship canal near Portage Bay, Se- attle, elev. 6 m, 7 Nov 2000, Zika 15655 (WTU). Previous knowledge. Willow-leaved cotoneaster is na- tive to western China, and planted as an ornamental in western Washington for its brilliant autumn fruits. Amer- ican robins (Turdus migratorius) and American crows (Corvus brachyrhynchos) eat the fruit and disperse the seed. 196 Significance. First collections for Washington as an es- cape from cultivation. COTONEASTER TENGYUEHENSIS J. Fryer & B. Hylm6 (RO- SACEAE).—King Co., thickets near Washington Park, Seattle, elev. 35 m, 15 Sep 2000, Zika 15482 (WTU); slope near small creek, Washington Park arboretum, Se- attle, elev. 20 m, 2 Nov 2000, Zika 15630 (WTU). Previous knowledge. Tengyueh cotoneaster is native to SW China, and an uncommon ornamental in western Washington. Significance. First report of an escape from cultivation in Washington. CREPIS SETOSA Haller f. (ASTERACEAE).—Clark Co., grassy roadside near Loop Road, elev. 10 m, 24 Jul 2000, Zika I5114A & Weinmann (WTU); lawn weed by soccer field, NE18th St., elev. 90 m, 14 Sep 2001, Zika 16558 (WTU). Significance. First collections in Washington for this southern European native. CYPERUS DIFFORMIS L. (CYPERACEAE).—Franklin Co., sandy E shore of free-flowing Columbia River, elev. 100 m, TION R28E S1, 2 Oct 2001, Zika 16671 (EIU, MICH, WS, WTU). Significance. First report for Washington for this Asian native. FRAXINUS PENNSYLVANICA Marsh. (OQLEACEAE).—Grant Co., low ground between dunes, near Potholes Wildlife Area, SW of Moses Lake, elev. ca. 330 m, 14 Jun 2001, Zika 16258 (WTU); King Co., wet thicket, Madrona Park, W shore of Lake Washington, Madrona, Seattle, elev. 5 m, 2 Aug 2000, Zika 15184 (WTU). Previous knowledge. Green ash is native to eastern North America, west to Montana, and planted as an or- namental in the Pacific Northwest. Significance. First record for Washington escaping from cultivation and naturalizing. GALIUM PEDMONTANUM (Bellardi) All. (RUBI- ACEAE).—Klickitat Co., disturbed meadow, Conboy Na- tional Wildlife Refuge, elev. 570 m, 15 Jun 2001, Rodman 508 et al. (WTU). Significance. First report for Washington for this Eu- ropean native. GERANIUM PYRENAICUM Burm. f. (GERANIACEAE).— King Co., gravel alleys and waste ground, Madrona, Se- attle, elev. 95 m, 14 May 2000, Zika 14969 (OSC, WS, WTU). Previous knowledge. Hedgerow cranesbill is native to Europe, grown in gardens, and known as a weed in Cal- ifornia and eastern North America. Significance. First report in Washington as an escape from cultivation. HIERACIUM LACHENALII C.C. Gmel. (ASTERACEAE).— King Co., sunny roadside, Route 410, Greenwater, 14 Jun 2001, Walker s.n. (WTU); Skamania Co., north ridge of Table Mountain, 25 Jun 2000, Arnett s.n. (WTU); Sno- homish Co., logged area, Perry Creek trail, 5 Aug 1962, Kruckeberg 5515 (WTU); common on roadside, Route 2 east of Index, elev. 300 m, 5 Jun 2000, Zika 15095d (WTU). Previous knowledge. Hieracium lachenalii s. str. (syn. H. acuminatum Jord.) is native to Europe and adventive MADRONO [Vol. 49 in eastern North America. All reports of H. vulgatum Fries from Washington are H. lachenalii, except one collection of true H. vulgatum from Pacific Co. (Maxwell 215 WTU). Significance. First collections for Washington. HIERACIUM MURORUM L. (ASTERACEAE ).—Pierce Co.., roadside and adjacent forest, Route 706 at Westside Road, Mt. Rainier National Park, elev. 640 m, 14 Aug 1999, Biek 2 (WTU). Previous knowledge. Wall hawkweed is native to Eu- rope, and adventive in eastern North America. Reports of H. atratum Fries from Washington belong here. Hieracium murorum has been collected as a weed in Portland, Oregon (Ornduff 6196 OSC, WTU). Significance. First collection for Washington. HIERACIUM SABAUDUM L. (ASTERACEAE ).—King Co., roadside, Interstate 90, 16 km E of North Bend, elev. 420 m, 20 Sep 2001, Brunskill s.n. (WTU); Skagit Co., road- side, Cain Lake Road near Alger, 3 Sep 1996, Lantz s.n. (WTU); Whatcom Co., E Lake Samish Road, ca. 10 km S of Bellingham, 2 Sep 1990, Burnett 280 (WTU); Inter- state-5, near S end of Samish Lake, elev. 30 m, 9 Sep 2000, Zika 15465 (WTU). Previous knowledge. Savoy hawkweed is native to Eu- rope and adventive in eastern North America and British Columbia. Reports of H. laevigatum Willd. from Wash- ington belong here. Significance. First report for Washington. HYPERICUM MACULATUM Crantz subsp. OBTUSIUSCULUM (Tourlet) Hayek (CLUSIACEAE).—King Co., crack in concrete sidewalk, Montlake, Seattle, elev. 25 m, 22 Jul 2001, Zika 16393 (WTU). Previous knowledge. Dotted St. Johnswort is native to Europe, and adventive in southern British Columbia. In Seattle it is spreading from an introduction in a “wild- flower”’ seed mix. Significance. First report for Washington. JUNCUS PATENS E. Mey. (SUNCACEAE).—Clark Co., shade of Fraxinus, Lackamas Creek floodplain, elev. ca. 65 m, 22 Mar 2001, Zika 15799 (WTU); low ground near SE Ist Street, Grass Valley, elev. ca. 70 m, 22 Mar 2001, Zika 15802 (WTU). Previous knowledge. Native in the Willamette Valley of Oregon, 20 km to the S. “Reported but not seen from Washington”’ (Hitchcock, Cronquist, and Ownbey, 1969, Vascular Plants of the Pacific Northwest, Part 1, Univ. of Washington Press). Significance. First collections to document this native in Washington. MOENCHIA ERECTA (L.) P. Gaertn., B. Mey. & Scherb. (CARYOPHYLLACEAE).—Pierce Co., dry prairie rem- nant, with Lepidium heterophyllum, N of Muck Creek, elev. ca. 120 m, 4 Jun 2001, Zika 16157 & Weinmann (WTU); dry prairie remnant, Route 507, 5 miles NE of Roy, elev. ca. 120 m, 4 Jun 2001, Zika 16166 & Wein- mann (WTU). Previous knowledge. Upright chickweed is native to Europe, and adventive in Oregon and British Columbia. Significance. First report for Washington. PHOTINIA DAVIDIANA (Decne.) Cardot (ROSACEAE).— King Co., bird-sown epiphyte in tree, near Lake Washing- ton, Martha Washington Park, Seattle, elev. 10 m, 6 Jun 2001, Zika 16184 & Jacobson (UBC, WTU); Kitsap Co., 2002] with Alnus rubra, pondshore, Bloedell Reserve, N end of Bainbridge Island, Puget Sound, elev. 30 m, 15 Nov 1999, Zika 14724 & Jacobson (WTUV). Significance. First collections of garden escapes for this native of China. PHOTINIA VILLOSA (Thunb.) DC. (ROSACEAE).—King Co., moist ground, with Alnus rubra, Rubus spectabilis, Union Bay, Seattle, elev. 5 m, 29 Sep 2000, Zika 15524 (WTU); rare adventive, Volunteer Park, Seattle, elev. 130 m, 8 Nov 2000, Zika 15670 & Jacobson (WTU). Previous knowledge. Oriental redtip is native to E Asia, and known as a garden escape in the eastern United States. Both King Co. sites are adjacent to ornamental plantings, and the species was apparently spread by frugivorous birds. Significance. First report from Washington as an escape from cultivation. PYRACANTHA COCCINEA M. Roem. (ROSACEAE).—King NOTEWORTHY COLLECTIONS 197 Co., near shore of Portage Bay, Seattle, elev. 5 m, 27 Aug 1999, Zika 14136 & Jacobson (WTU); San Juan Co., 3.5 km SE of Sportsman Lake, San Juan Island, 24 Oct 1999, Zika 14641 (WTU). Significance. First Washington report as an escape from cultivation. STACHYS ARVENSIS (L.) L. (LAMIACEAE).—King Co., weed on gravel roadside, near S shore of Steele Lake, Federal Way, 3 Apr 2001, Zika 15866 (WTU). Significance. First collection in Washington for this Eu- ropean native. VERONICA VERNA L. (SCROPHULARIACEAE).—Asotin Co., Route 129, Buford Cr., elev. 600 m, 30 May 1991, Zika 11135 (OSC, WTU, WS); Chelan Co., N shore Lake Chelan, elev. 335 m, 7 Jun 1998, Zika 13427 (WTU). Significance. First report for Washington. —PETER FE ZiIkA, Herbarium, Dept. of Botany, Box 355325, Univ. of Washington, Seattle, WA 98195-5325. MADRONO, Vol. 49, No. 3, p. 198, 2002 REVIEW A cactus odyssey: Journeys in the wilds of Bolivia, Argentina, and Peru. By JAMES D. MAUSETH, ROB- ERTO KIESLING, AND CARLOS OSTOLAZA. 2002. Tim- ber Press, Portland, OR. 306 pp. ISBN 0-88192- 526-8. This wonderful book presents an engrossing ac- count of the authors’ botanical fieldwork in all manner of habitats throughout much of South America Over a seven-year period. The relaxed nar- rative style, beautiful photographs, and liberal in- terjection of humor make this an entertaining read for just about anyone. Indeed, approachability by the layperson is a stated goal of the work, and one that is met very admirably. For botanists, the book will be even more absorbing, as intriguing aspects of taxonomy, morphology, anatomy, ecology and physiology of cacti are highlighted throughout. As a cactus freak, I found the work completely en- thralling. I am very inspired by the authors’ philosophy, stated clearly at the beginning and throughout the work: that cooperation, openness and sharing of findings among biologists is beneficial to all. It is quite refreshing to view this opinion in print. The fruits of this philosophy are evident in the quality and number of publications resulting from the field- work documented here (Kielsing 1995; Mauseth and Kielsing 1997; Ostolaza 1997; Mauseth 1999, 2000; among others). Seven chapters are included: one on cactus bi- ology, and two each on the fieldwork conducted in Bolivia, Peru, and Argentina, respectively. The in- troduction to cacti is broadly written and accessible, resulting in a concise yet thoughtful description of the family, its evolution, ecology, and taxonomy. My one minor criticism of the book is in this intro- duction, where the movement of an ocean current is described somewhat inaccurately (p. 22). The cir- cumglobal southern ocean current (Bartholomew 1958, Plate 2) precludes movement of water from the Atlantic to the Pacific between South America and Antarctica. This is the most minor of criticisms, however. Subsequent chapters entwine absorbing vignettes of all manner of cacti in a chronological framework of field experience. The discovery of each new tax- on on the journey is used to highlight one or more interesting facets of cactus biology. For example, the appearance of Melocactus includes a discussion of cephalia, or the sight of Prosopis (Fabaceae) and Prosopanche (Hydnoraceae) touches off a discus- sion of mistletoes found in cacti. The authors con- sistently relate the spotlighted theme for that taxon to other plants, often drawing parallels between the plants encountered and other cacti more familiar to the North American reader. The authors have done a tremendous job here; I cannot emphasize enough how captivating and diverse a portrait of cacti is presented in this way. The biological material pre- sented is written accessibly, but without apparent oversimplification. I also appreciate the time spent describing those aspects of cactus biology that are not easily understood. The authors’ repeated ap- peals for students to study these phenomena are a welcome incorporation in the work. Equally appealing is the description of the va- garies of the field. The less-than-ideal road condi- tions, rough accommodations, cold food, unsym- pathetic authorities and vehicle breakdowns of fieldwork are related with the dry wit that I have associated with Mauseth since taking his plant anat- omy course in 1997. Here, his sense of humor is unerring. Anyone who has spent time afield will find themselves at least smiling but probably laugh- ing outright at the dead-to-rights depiction of being constantly outpaced by the sun. The photographs are all excellent, and well cho- sen to illustrate particular points. South America is a beautiful, wild, diverse place as pictured here, and the cacti are flat-out gorgeous. My own favorites include figures of Browningia candelaris and Azur- eocereus. Many wonderful landscape and habitat shots are included, as well as several graphical de- pictions of life in the field. This book has something for everyone, and I rec- ommend it to all. I am in the middle of my second reading, and it has not lost an ounce of my interest. —M. PATRICK GRIFFITH, Rancho Santa Ana Botanic Gar- den, 1500 N. College Avenue, Claremont, CA 91711. E-mail: michael.patrick.griffith @cgu.edu. LITERATURE CITED BARTHOLOMEW, J. 1958. The Times atlas of the world: mid-century edition, Vol. 1. Times Publishing Com- pany, London, U.K. KIELSING, R. 1995. Argentine notocacti of the genus Par- odia. Cactus and Succulent Journal (U.S.) 67:14—22. MAUuSETH, J. D. 1999. Comparative anatomy of Espostoa, Pseudoespostoa, Thrixanthocereus, and Vatricania (Cactaceae). Bradleya 17:33—43. MAusETH, J. D. 2000. Theoretical aspects of surface-to- volume ratios and water-storage capacities of succu- lent shoots. American Journal of Botany 87:1107— NS), MAUuSETH, J. D. AND R. KIELSING. 1997. Comparative anat- omy of Neoraimondia roseiflora and Neocardenasia herzogiana (Cactaceae). Haseltonia 5:37—S0. OSTOLAZA, C. 1997. Cactus del sur de Cajamarca y del valle del Rio Sana. Quepo 11:57—68. MADRONO, Vol. 49, No. 3, p. 199, 2002 ANNOUNCEMENT BIENNIAL GRADUATE STUDENT MEETING AND ANNUAL BANQUET 15 FEBRUARY 2003 AT THE UNIVERSITY OF SAN DIEGO The California Botanical Society’s Biennial Graduate Student Meeting and Annual Banquet will be held on Saturday, 15 February 2003 at the Hahn University Center of the University of San Diego. Graduate students everywhere who are initiating, conducting, or finishing research projects in any area of botany (e.g., ecology, evolution, conserva- tion, floristics, morphology, development, etc.) are encouraged to attend the meeting (and banquet) and to give short oral presentations on their research plans or findings. The venue is an ideal opportunity for students to gain experience giving presentations in the standard format of scientific meetings, to meet students involved in botanical research from other institutions, and to learn more about botany in general. Presentations will be judged by student peers and awards for best papers in proposed re- search, research-in-progress, and completed re- search will be presented at the evening banquet. Abstracts of all presentations will be published on- line at the California Botanical Society web-site (www.calbotsoc.org). Our speaker for the annual banquet will be Dr. Jon Rebman, Curator of the Herbarium at the San Diego Natural History Museum, who will present an after-dinner lecture entitled “‘Discoveries on a Floristic Frontier: Baja California.’’ Dr. Rebman’s botanical explorations of remote regions of Baja California are exemplary of binational collabora- tion between the US and Mexico and have yielded many exciting findings, including plants previously unknown to science and new insights into cactus biology and evolution. We look forward to a fas- cinating evening of highlights from Dr. Rebman’s field research in beautiful and rugged desert ranges, such as the Sierra de la Giganta, where he will be involved in a major expedition this fall. The graduate student meeting and annual ban- quet are open to CBS members and non-members alike; anyone interested in the meeting and/or ban- quet is encouraged to attend. Registration infor- mation for the meeting and banquet will be forth- coming. MADRONO, Vol. 49, No. 3, p. 200, 2002 ERRATUM STuTZ, H. C., M. R. STUTZ, AND S. C. SANDERSON. 2001. Atriplex robusta (Chenopodiaceae), a new perennial species from northwestern Utah. Madro- no 48:112—115. The name “Atriplex robusta’ H. C. Stutz, M. R. Stutz, & S. C. Sanderson (Madrono 48:112. 2001) is illegitimate, having already been used (A. robusta Speg. In Gand.) Atriplex tridentata Kuntze var. robusta H. C. Stutz, M. R. Stutz, & S. C. Sanderson, var. nov.—TYPE: USA, Utah, Tooele Co., 1 mi W of Knolls, T15 R13W S15, shoulder of highway I-80, 1280 m elevation, 16 Sep 1977, H. C. Stutz $141 (Holotype: BRY; Isotypes, BRY, CA, CAS, GH, MO, NY, RM, UC). Frutices caespitosi, 40—80 cm alti. Caules erecti vel ascendentes, ramosi a basi ad apicem, dense furfuraceus, 1-8 mm diam., fragilis. Folia oblonga, ascendentia usque appressa, dense furfuraceae; fo- lia ephemera verna et aestiva 15—30 mm longa, 5— 10 mm lata; folia serotina aestiva et hiberna 3—10 mm longa, 2—5 mm lata, anatomia foliaris Kranz- typi. Plantae dioeciae, raro monoeciae. Flores stam- inati sessiles, ad brevi-ramulus axillares in angusti paniculas terminales; perianthium campanulatum, 5-partitum ad medium, dense furfuraceum, segmen- tis ovatis usque ellipticis, 2 mm longis, | mm latis; stamina 5, filamentis | mm longis, antheris ca. 2 mm longis, | mm latis. Flores pistillati solitarii, ses- siles, in pleurumque sine foliis confertas paniculas terminales. Bracteae fructiferae furfuraceae, com- pressae, urceolatae, latissimae infra media, 5 mm latae, 7-8 mm longae, exappendiculatae, cum 3—10 marginalibus dentibus, 0.5—2 mm longis, qui me- dianus longissimus. Utriculus orbiculatus, pericar- pio membranceo pellucido. Semena 5 mm diam., testa membranacea, brunnea; radicula supera. Volume 49, Number 3, pages 137—200, published 17 December 2002. SUBSCRIPTIONS—MEMBERSHIP Membership in the California Botanical Society is open to individuals ($27 per year; family $30 per year; emeritus $17 per year; students $17 per year for a maximum of 7 years). Late fees may be assessed. Members of the Society receive MAproNo free. Institutional subscriptions to MApDRONO are available ($60). Membership is based on a calendar year only. Life memberships are $540. Applications for membership (including dues), orders for sub- scriptions, and renewal payments should be sent to the Treasurer. 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Institutional abbreviations in specimen citations should follow Holmgren, Keuken, and Schofield, Index Herbariorum, 8th ed. Names of authors of scientific names should be abbreviated according to Brummitt and Powell, Authors of Plant Names (1992) and, if not included in this index, spelled out in full. Titles of all periodicals, serials, and books should be given in full. Books should include the place and date of publication, publisher, and edition, if other than the first. All members of the California Botanical Society are allotted 5 free pages per volume in MApRoNo. Joint authors may split the full page number. Beyond that number of pages a required editorial fee of $40 per page will be assessed. The purpose of this fee is not to pay directly for the costs of publishing any particular paper, but rather to allow the Society to continue publishing MapRONo on a reasonable schedule, with equity among all members for access to its pages. Printer’s fees for illustrations and typographically difficult material @ $35 per page (if their sum exceeds 30 percent of the paper) and for author’s changes after typesetting @ $4.50 per line will be charged to authors. At the time of submission, authors must provide information describing the extent to which data in the manu- script have been used in other papers that are published, in press, submitted, or soon to be submitted elsewhere. jf M a5 é VOLUME 49, NUMBER 4 OCTOBER-—DECEMBER 2002 MADRONO A WEST AMERICAN JOURNAL OF | IND va SN Wis ai i wlll | EW al | CONTENTS FLOWERING PATTERNS AND REPRODUCTIVE ECOLOGY OF MAMMILLARIA GRAHAMII (CACTACEAE), A COMMON, SMALL CACTUS IN THE SONORAN DESERT HOT CB LEST ON ON RTI Aa aes Re Pr Ra Oe ne 201 THE EFFECT OF FIRE AND COLD TREATMENTS ON SEED GERMINATION OF ANNUAL AND PERENNIAL POPULATIONS OF ESCHSCHOLZIA CALIFORNICA (PAPAVERACEAE) IN SOUTHERN CALIFORNIA Arlee M. Montalvo, Laura J. Feist-Alvey, and Catherine E. Koehler ...... 207 TEMPERATURE LIMITATIONS FOR CULTIVATION OF EDIBLE CACTI IN CALIFORNIA Park S. Nobel, Erick De la Barrera, David W. Beilman, Jennifer H. ID OMCTIN: GING Mate ONG Is ZU essa athe SP ea con a ees os ad sc ooaggpsaaasaseasedendes 228 Six New SPECIES AND TAXONOMIC REVISIONS IN MEXICAN GAUDICHAUDIA (MALPIGHIACEAE) SUCVEI ME VCSSUD r,s Sence nce dscneck saeco see Me uae Op ao soon eye UR Pn hnn sehen vaeindeauese 237 RETICULATE ANCESTRY IN MEXICAN GAUDICHAUDIA (MALPIGHIACEAE) ANALYZED WITH RAPD's AND SOUTHERN HYBRIDIZATION Steven L. Jessup’ .... 2075 IY. ACE s AZ LAPS oy aev en cnnserensnnenaseee 256 LONG-TERM POPULATION DyNAMICS OF NATIVE NASSELLA (POACEAE) BUNCHGRASSES IN CENTRAL CALIFORNIA Jason G. Hamilton, James R. Griffin, and Mark R. Stromberg ................ 274 A NEw SPECIES OF PRUNUS (ROSACEAE) FROM THE MOJAVE DESERT OF CALIFORNIA BREA IO A EAU ME A Nae ses MY sn one MUN Sea A eee ove ve ve ntneneceoeeee 285 A NEw CEANOTHUS (RHAMNACEAE) SPECIES FROM NORTHERN BAJA CALIFORNIA, MEeExIco Steve Boyd anal dor FE. Keeley! iso ON. cascccnes oe p ew wae evenseesseceeseserenes 289 COLLINSIA ANTONINA IS EVOLUTIONARILY DISTINCT FROM C. PARRYI (SCROPHULARIACEAE SENSU LATO) BriceG, Baldwin and W. SCOtt AVRIBTUSIET ......<...0cssssoesecsvesecseeveeeseevsseoues 295 BOOK REVIEW FIELD GUIDE TO LIVERWORT GENERA OF PACIFIC NORTH AMERICA, BY W.B. SCHOFIELD Sa Eat SN TN OGG eh tee a ee eae dot a sleds vesteSouW aeiGe nab vbsvniade Usavenvenies 298 ANNOUNCEMENTS Perse GREP ORE BOR VOLUME A 2 occ caevesessuvevsetnccavcnctutancunsiecncersssevreecseccneeses 299 BNE, SEP CRUE EUR) VOW LINID AO ..5.cccccbesssccUel oeetsesnctudtonviwavsdéuccsiuvadeveosvavenvesesesseoenens 300 ESTE TEES STEED 1 27 SO i ee 301 I egien Sam N NDEI ie ys as gsc Seo un Cee cen cu cideavevovanrecbuvedkadddussvosUviversessvvesneseee 302 ie abate MEAT TPE OY Feet APIA i, 2. A coescchcescovesoceendenatevncndaeusessvivesewssssecceusnosseete 305 Par EE OBE INES HOR. V OLLIE A oo oo acc cso reakeeneeaceovactsscesivsserootcssecesalsnccsoonecees il aE Sa MANU PRN Esc tee icc tac neva caw oc cusevncecsesnvonewvni es dovenevisdveersbneeeen- iV PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY Maprono (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. Postmaster: Send address changes to MApRONO, Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. Editor—Dr. JOHN CALLAWAY Dept. of Environmental Science University of San Francisco 2130 Fulton Street San Francisco, CA 94117-1080 callaway @usfca.edu Book Editor—Jon E. KEELEY Noteworthy Collections Editors—DiETER WILKEN, MARGRIET WETHERWAX Board of Editors Class of: 2002—-NorMaAN ELLSTRAND, University of California, Riverside, CA Cara M. D’ Antonio, University of California, Berkeley, CA 2003—-FREDERICK ZECHMAN, California State University, Fresno, CA Jon E. KeELEY, U.S. Geological Service, Biological Resources Division, Three Rivers, CA 2004—Davip M. Woon, California State University, Chico, CA INGRID Parker, University of California, Santa Cruz, CA 2005—J. Mark Porter, Rancho Santa Ana Botanic Garden, Claremont, CA Jon P. REBMAN, San Diego Natural History Museum, San Diego, CA CALIFORNIA BOTANICAL SOCIETY, INC. OFFICERS FOR 2002—2003 President: BRUCE BALDwin, Jepson Herbarium and Dept. of Integrative Biology, 1001 Valley Life Sciences Bldg. #2465, University of California, Berkeley, CA 94720. First Vice President: Rov Myatt, San José State University, Dept. of Biol. Sciences, One Washington Square, San José, CA 95192. rmyatt@email.sjsu.edu Second Vice President: MicHaeL S. Mayer, Department of Biology, University of San Diego, San Diego, CA 92110, mayer @sandiego.edu Recording Secretary: Stact Markos, Friends of the Jepson Herbarium, University of California, Berkeley, CA 94720- 2465, smarkos @ socrates.berkeley.edu. Corresponding Secretary: SUSAN BAINBRIDGE, Jepson Herbarium, 1001 VLSB #2465, University of California, Berkeley, CA 94720-2465. 510/643-7008. suebain@SSCL.berkeley.edu Treasurer: Roy Buck, % University Herbarium, University of California, Berkeley, CA 94720. The Council of the California Botanical Society comprises the officers listed above plus the immediate Past President, R. Jonn LittLe, Sycamore Environmental Consultants, 6355 Riverside Blvd., Suite C, Sacramento, CA 95831; the Editor of Maprono; three elected Council Members: James SHEVOCK, National Park Service, 1111 Jackson St., Suite 700, Oakland, CA 94607-4807. 510/817-1231; ANNE BRADLEY, USDA Forest Service, Pacific Southwest Region, 1323 Club Drive, Vallejo, CA 94592. abradley @fs.fed.us; DEAN KELcH, Jepson and University Herbarium, University of California, Berkeley, CA 94720, dkelch @sscl.berkeley.edu. Graduate Student Representative: ELIZABETH ZACHARIAS, Department of Integrative Biology, University of California, Berkeley, CA 94720. ezachar @socrates.berkeley.edu; Local (San Diego) Graduate Student Representative: RoBert K. Lauri, Department of Biology, San Diego State Uni- versity, San Diego, CA 92182. Webmasters: J. Curtis CLarK, Department of Biological Sciences, California State Polytechnic University, Pomona, CA 91768; Joun C. LADuke, Department of Biology, University of North Dakota, Grand Forks, ND 58202. This paper meets the requirements of ANSI/NISO.Z39.48-1992 (Permanence of Paper). MabpDRONO, Vol. 49, No. 4, pp. 201—206, 2002 FLOWERING PATTERNS AND REPRODUCTIVE ECOLOGY OF MAMMILLARIA GRAHAMII (CACTACEAE), A COMMON, SMALL CACTUS IN THE SONORAN DESERT JANICE E. BOWERS U.S. Geological Survey, 1675 West Anklam Road, Tucson, AZ 85745 jebowers @usgs.gov ABSTRACT Mammillaria grahamii is an outcrossing species that can flower as many as four times a year. The number of flowers produced by an individual plant depends largely on plant volume. Fruit set is high, about 77%. Flowers normally live one day but can open a second day when pollination is inadequate. Flower production within a population varies interannually and increases with rain in the week before flowering. Even heavy rains do not induce all potentially fertile tubercules to flower during a single event. Because some tubercules are withheld, plants can flower several times a year, given appropriate conditions. Mammillaria grahamii plants risk poor fruit set if they happen to flower when pollinator populations are low or when pollinators preferentially visit other species. Withholding of tubercules might allow plants to spread this risk across the entire flowering season, thus increasing the probability that flowering will coincide with pollinator activity. Key Words: Cactaceae, floral biology, floral longevity, Mammillaria grahamii. When and how often plants bloom has profound consequences for seed set, seed dispersal, and seed- ling recruitment, thus making the study of flower- ing phenology an important aspect of plant popu- lation biology (Primack 1985; Rathcke and Lacey 1985; Newstrom et al. 1994). Because flowering patterns affect the behavior and reproductive suc- cess of pollinators and granivores, phenology un- derlies the study of evolutionary and community ecology as well (Brody 1997). Knowledge of flow- ering patterns is, moreover, crucial to conservation and management of rare species (Newstrom et al. 1994). For example, Coryphantha_ robustispina (Schott ex Engelmann) Britton & Rose subsp. scheeri (Muehlenpfordt) N. P. Taylor, a small, en- dangered cactus, can bloom several times per year, but because the proportion of sexually mature plants that flower during any single event varies considerably (Roller 1996), within-year flowering patterns must be considered when estimating repro- ductive effort. In general, flowering patterns and reproductive biology of small cacti in the arid southwestern United States are poorly known (Johnson 1992). The few species that have been studied are for the most part endangered or threatened, giving us a poor idea of what phenological and floral traits are associated with successful reproduction and estab- lishment. This paper examines the reproductive ecology of Mammillaria grahamii Engelmann, a small cactus that is common and widespread in and around the Sonoran Desert (Shreve and Wiggins 1964; Aguilar et al. 2000). The species has been known as M. microcarpa Engelmann, not a validly published name (Aguilar et al. 2000). The reproductive ecology of M. grahamii has not been studied in any detail. Flowers are bowl-shaped and are relatively small for the family, about 2 to 4 cm in diameter (Anderson 2001). They arise from areoles located in the axils of old tubercules, that is, tubercules formed the previous year or earlier (Shreve and Wiggins 1964). Tubercules are modi- fied leaf bases that appear as regularly spaced pro- jections on the plant body. Once an areole has flow- ered, it cannot flower again (Gibson and Nobel 1986). Petals are deep pink or white with a pink midrib. Size, shape, and diurnal habit of the flowers indicate that they are bee-pollinated (Grant and Grant 1979). Stems are solitary or branching and grow to a height of 10 to 20 cm. Maximum lifespan is about 11 years (Goldberg and Turner 1986). Mammillaria grahamii can flower several times per year and in any month from March to Septem- ber (personal observation; M. Dimmitt personal communication). The ultimate goal of this study was to determine the benefits of multiple flowering events and the conditions under which they occur. Specific objectives were to determine: 1) relation between plant size and flower production, 2) effect of rain on flower production, 3) longevity of indi- vidual flowers, 4) minimum reproductive size, 5) breeding system, and 6) fruit set and seed produc- tion. METHODS Study area. The study site is located at 720 m above sea level about 11 km northwest of Tucson, Pima County, Arizona (32°16'N, 111°2’W). Terrain is a level to gently sloping alluvial terrace. Soils are derived from rhyolitic parent material, and the surface is gravelly to cobbly. Vegetation is char- acteristic of the Arizona Upland subdivision of the 202 Sonoran Desert (Shreve and Wiggins 1964). Dom- inant plants include Cercidium microphyllum (Torr.) Rose & Johnston, Ambrosia deltoidea (A. Gray) Payne, Acacia constricta Benth., Larrea tri- dentata (Moc. & Ses.) Cav., Krameria grayi Rose & Painter, Carnegiea gigantea (Engelmann) Britton & Rose, Opuntia engelmannii Salm-Dyck., Fero- cactus wislizeni (Engelmann) Britton & Rose and Opuntia leptocaulis DC. Annual precipitation (300 mm) is seasonally distributed as a highly variable winter and early spring (November to March), an arid late spring (April to June), a predictable sum- mer monsoon (July to August), and a highly vari- able autumn (September to October). Maximum temperatures in summer often exceed 40°C. Mini- mum temperatures rarely drop below —6°C in win- ter. Although freezing nights can be frequent in winter, daytime temperatures always rise above O°C. Flower production and plant size. On July 24, 1996, height and diameter of all stems on 60 M™. grahamii plants were measured to the nearest 0.5 cm, and the number of flowers and flower buds on each stem was counted. Plants too small to flower were excluded from the sample. Stem volume was approximated using the formula for a cylinder, then individual volumes were summed to get volume of the entire plant. Number of flowers was used as the dependent variable in separate linear regressions against height of the tallest stem, diameter of the thickest stem, number of stems, and plant volume. Annual flowering patterns. For monitoring fre- quency, duration, and intensity of bloom, all M. grahami plants (n = 68) within an area approxi- mately 15 m by 20 m were marked on July 27, 1996 by affixing numbered aluminum tags to the ground next to the plant. The sample included some but not all of the plants sampled previously and also included plants that were too small to flower. Height and diameter of the tallest stem of each plant were measured to the nearest 0.5 cm. The number of open flowers on each plant in the sample was counted daily from the first to the last day of bloom whenever the sample flowered in 1996 (with one exception, described below) and 1997. In ad- dition, open and spent flowers were counted on the last day of a single blooming event in 1999. Daily values in 1996 and 1997 were summed to deter- mine the total number of flowers produced per plant during each event. Flower production during the first blooming event of 1996 was determined on the last day of the event by counting number of open and spent flowers. At the first event in 1996, flow- ers were counted on 25 single-stemmed plants only. Thereafter, all marked plants were sampled at every event. Rain and flower production. The effect of rain on flower production was assessed for six flowering events in 1996, 1997, and 1999 using only those plants for which flowers had been counted at all six MADRONO [Vol. 49 events (n = 15 plants). The mean number of flow- ers per plant at each event was calculated, then Spearman rank-order correlation was used to deter- mine the strength of association between flower production and rain (mm) in the weeks before flow- ering. Pollinator visitation. Visits by potential pollina- tors were monitored from August 3 to 6, 1996. Al- together, 25 flowers on 14 plants were watched for 10 minutes each, during which time the number of bee visitors was recorded. No attempt was made to identify the bees. Beetles, which are not effective pollinators of cactus flowers (Grant and Connell 1979), were not included in these surveys. Obser- vations were made between 0830 and 1130 hr, the period of greatest pollinator activity. Characteristics of fruits and seeds. Twenty-four ripe fruits were collected in September 1996 and individually weighed to the nearest 0.001 g. Length and width of each fruit were measured to the near- est 0.5 mm. Seeds were removed and air-dried, then the mass of seeds from each fruit was measured to the nearest 0.001 g. Seed set was determined by counting the number of seeds in each fruit. The weight of an individual seed was calculated by di- viding seed mass by number of seeds. Fruit set. Fruit set, defined as the proportion of flowers that produced fruits, was studied during two separate blooming events in August 1997. Thirty reproductively mature plants, none included in the previous samples, were numbered and tagged. Fif- teen were randomly selected and covered by wire mesh cages to prevent pollinators from getting ac- cess to the flowers. The number of set fruits was counted several weeks after flowering ended. Sep- arate Mann-Whitney tests were used to determine the effect of treatment (caged or open-pollinated) and blooming event (first or second) on the pro- portion of flowers that set fruit. Flower longevity. The lifespan of individual flowers was studied on the same sample of 30 plants during the first summer blooming event in August 1997. All flowers opening for the first time were marked on a daily basis with glass-headed pins, using a different color of pin for each day. The number of freshly opened flowers and the num- ber that opened more than once were counted on each plant every day. Breeding system. Pollination and germination ex- periments in a greenhouse were used to determine the breeding system. Altogether, 10 self pollinations (two flowers from the same plant) and 6 cross pol- linations (two flowers from different plants) were made. For each cross, one of the flowers was tagged with a label indicating date and cross num- ber. Stamens were removed from both flowers, then the stamens of the untagged flower were used to pollinate the stigmas of the tagged flower. Tagged 2002] iz ® = 2 1997 TgeOut 1 60 + =| 40 - - 20 | =| : L 146 166 186 206 226 246 266 Day of Year Fic. 1. Flowering curves for Mammillaria grahamii in 1996 (top) and 1997 (bottom). Y axis represents percent of sample in flower; n = 68 except for first event of 1996, where n = 25. The first event in 1996 lasted five days but flowers were counted only on the final day. flowers were checked regularly for developing fruits. Mature fruits were collected and dried, and the number of seeds in each was counted. Seed germination was tested using fruits from cross and self pollinations. Twenty seeds from each fruit were planted in a four-inch-square plastic pot on a moistened, sterilized mixture of sand, turface, and milled peat moss. The pots were covered with clear plastic wrap to prevent desiccation while al- lowing exposure to light. After two months, the covers were removed and the pots watered weekly during the next three months, then every two weeks for the following seven months. The number of liv- ing and dead seedlings in each pot were counted at two months and twelve months. RESULTS Annual flowering patterns. As reported previous- ly, M. grahamii can flower three to four times in a year (Fig. 1). Flower dates in this study were as follows: July 21 to 25, 1996; August 2 to 11, 1996; May 26 to 30, 1997; August 14 to 20, 1997; August 29 to September 3, 1997; and September 21 to 23, 1997. In both years, the most intense blooming events occurred in summer and lasted more than a week (Fig. 1). Typically, 60 to 80% of the sample flowered during the first summer event. Later sum- mer events involved fewer plants and lasted only three to four days. No flowers were produced in spring of 1996. Spring flowering in 1997 involved about 40% of the population and lasted five days. Flower production and plant size. In the sample of reproductively mature plants (n = 60), volume BOWERS: FLOWERING PATTERNS OF MAMMILLARIA 203 100 — re ” 80 + = 10) = ® e = 60} re ro) ® = 40 ¢ ® e as 3 we ° y = 4.28 + 0.04x 20 ope ia e r = 0.75 o f= 4 cm in height produced flow- ers. Rain and flower production. Spearman correla- tion analysis showed that rain in the week before flowering was strongly correlated with mean num- ber of flowers produced by 15 plants during six flowering events (r, = 0.94, P < 0.05) (Fig. 3). 14 - = 42 - : oO a O Qa 2 [- T = (00) = LL ne) ~ = [ : oO ) E 2 = iS | | 3 = 5 Event Fic. 3. Correlation between flower production and rain- fall. Top: mean number of flowers/plant (+ 1 SE) at six flowering events. Bottom: rain (mm) in the week before flowering. First day of flowering as follows: event 1, Au- gust 2, 1996; event 2, August 21, 1996; event 3, May 27, 1997; event 4, August 15, 1997; event 5, August 29, 1997; event 6, July 14, 1999. 204 TABLE 1. FRUIT AND SEED CHARACTERISTICS, MAMMILLARIA GRAHAMII; PEARSON CORRELATION COEFFICIENTS (N = 24 Fruits). * = Significant at P < 0.001. Fruit Fruit Fruit Seed mass length width mass Fruit length 0.54 Fruit width OFS 0.13 Seed mass 0.40 =().37 0.75% Seed number 0.45 =()),25 OVBe 0.89* Mean flower production was also correlated with rain in the two weeks before flowering (r, = 0.90, P < 0.05) but not with rain in the three weeks be- fore flowering (r, = 0.77, P > 0.05). Pollinator visitation. Visitation varied consider- ably over four days: visits/flower/hr (+SD) on the first through fourth days of observation were 0.0, 5.3 = 3.6, 5.0 = 2.4, and 41.1 + 25.3, respectively. Cloudy, humid weather on the first day of obser- vation may have limited pollinator activity; the re- maining days were clear and hot. On the second and third days of observation, all flower visitors were small bees, whereas on the fourth day, most visitors were honeybees. Characteristics of fruits and seeds. In the wild population, the number of seeds per fruit averaged 149 = 45 (SD). Fruit length and width (+SD) were, respectively, 26.0 mm + 5.2 mm and 7.2 mm + 0.9 mm. Mass of all seeds in a single fruit averaged 0.038 g + 0.016 g (SD). Calculated mass of an individual seed was 0.0003 g. Seed mass constitut- ed on average 9.1% of fresh fruit mass. Fruit width was the best predictor of number of seeds, seed mass, and fruit mass (Table 1). In addition, seed mass and number of seeds were highly correlated (Table 1). Fruit set. Fruit set of caged and open-pollinated plants averaged, respectively, 1.4% and 68.0% in the first blooming event, 1.1% and 85.0% in the second. The difference between treatments was highly significant (Mann-Whitney U = 0.0, P = 0.001), but the difference between blooming events was not (Mann-Whitney U = 350.0, P = 0.70). Flower longevity. Most flowers, whether caged or not, opened only a single day. Nevertheless, the proportion of flowers that opened a second day was almost 14 times greater for caged (63 of 284 flow- ers) than uncaged plants (3 of 188 flowers). It ap- pears that flowers typically live a single day if the level of pollination is adequate and that they can sometimes open a second day if it is not. Breeding system. Fruit set was 100% for crossed flowers (6 of 6) but only 10% for selfed flowers (1 of 10). The single fruit produced from the self-pol- linated flowers contained 281 seeds, but none ger- minated. Thus, in the rare instance when self pol- lination yielded fruit, the seeds apparently were not MADRONO [Vol. 49 viable. The average number of seeds in fruits pro- duced by cross pollination was 208 + 45 (SD). This was considerably higher than in the wild population and probably reflects a difference in pollen loads. Seeds from cross-pollinated fruits were highly ger- minable. Two months after planting, germination averaged 11.7%. After twelve months, average ger- mination was 78.3%. DISCUSSION In some respects, the floral biology of M. gra- hamii is similar to that of other small cacti in the arid southwestern United States (Table 2). As for Echinomastus erectocentrus (J. M. Coulter) Britton & Rose (Johnson 1992) and Escobaria robbinso- rum (W. H. Earle) D. R. Hunt (Schmalzel et al. 1995), plants reach reproductive maturity at a small size, and flower production increases as plants grow. As with Sclerocactus polyancistrus (Engel- mann & Bigelow) Britton & Rose (May 1994), var- iability in flower production from year to year or event to event is strongly correlated with rain. In other respects, the reproductive strategy of M. grahamii, a common and relatively widespread spe- cies, differs from that of small cacti that are rare or narrowly distributed. The number of flowers per plant and seeds per fruit is considerably higher (Ta- ble 2). The potential flowering season is longer, as well, and there can be two to four times as many flowering events per year (Table 2), a pattern that Newstrom et al. (1994) describe as “‘subannual.”’ The abundance and distribution of any species nec- essarily arises from multiple intrinsic, environmen- tal, and historical factors and cannot be ascribed to reproductive biology alone. Nevertheless, the com- bination of subannual blooming, large number of flowers per plant, high fruit set, and high seed pro- duction makes M. grahamii substantially more fe- cund than other small cacti (Table 2) and doubtless contributes to its success. Small cacti are at high risk of illicit collection (Bennett et al. 1986). During one study, for exam- ple, collectors illegally removed 31% of 324 M. grahamii and 44% of 9 M. thornberi Orcutt (Ben- nett et al. 1986). The latter species is considered vulnerable (Nabhan et al. 1989). Although the per- centages are roughly equivalent, the number of plants remaining—23 versus 5—is not. Especially for small cacti that have low fecundity, such dep- redations can represent a substantial portion of the reproductive capacity of the population. It remains to be seen whether M. grahamii, like other small cacti in the region, is obligately out- crossing. In greenhouse experiments, only 1 of 10 self-pollinated flowers set fruit. Because these flow- ers received ample pollen, low fruit set was likely a consequence of self incompatibility rather than inadequate transfer of pollen from stamens to stig- mas. None of the seeds from the selfed fruit ger- minated, in contrast to seeds from cross-pollinated 2002] BOWERS: FLOWERING PATTERNS OF MAMMILLARIA 205 TABLE 2. COMPARATIVE FLORAL BIOLOGY OF SELECTED SMALL CACTI. Species abbreviations and data sources as follows: CORY, Coryphantha robustispina subsp. scheeri (Roller 1996); ECHI, Echinomastus erectocentrus (Johnson 1992): ESCO, Escobaria robbinsorum (Schmalzel et al. 1995); MAMM, Mammillaria grahamii; SCLE, Sclerocactus polyan- cistrus (May 1994). Flower and stem sizes are from Anderson (2001). Other abbreviations: SS, single-stemmed: MS, multiple-stemmed; n.d., no data. Information on status is from http://arizonaes.fws.gov and http://www.cnps.org. CORY ECHI ESCO MAMM SCLE Flower diameter 5-7 4-5 1-2 2-4 5 Stem height (cm) 5-15 10—37 2-6 7-20 10—40 Stem diameter (cm) 5-9 7-12 2-6 7-11 5-9 Flowers/plant/yr (range) 1—23 1-16 1-7 1-136 1-6 Flowers/plant/yr (mean) iL n.d. 2 23 4 Height at first lowering (mm) 30 24 13 DS 30 Breeding system Outcrossing Outcrossing Outcrossing Outcrossing Outcrossing Fruit set (percent) 71 94 93 a n.d. Seeds/fruit 89 oF 20 149-209 120 Germination (percent) 89 n.d. n.d. 12-78 “low” Flower longevity (days) ] n.d. n.d. 1 (2) a7 Flowering season May-—Jul Mar—Apr Mar—Apr Mar—Sep Apr—May Flowering events/yr 1-3 1 1 1-4 | Duration of events (days) 1 Dag | n.d. 3-11 n.d. Lifespan (yr) <30 n.d. ilg/ 11 17 Plant morphology SS, MS SS SS SS, MS SS Status Endangered Candidate Threatened Not listed “Watch fruits, which germinated at a high rate. Taken to- ACKNOWLEDGMENTS gether, the pollination and germination experiments suggest that M. grahamii is obligately outcrossing; additional experimental work is needed to deter- mine whether this is indeed the case. Because M. grahamii produces only a single flower in each axil, flower production in one year is ultimately limited by the number of tubercules produced in previous years and by the proportion of old tubercules that have already flowered. An- nual tubercule production is in turn determined by plant volume and, probably, rainfall. The intensity of bloom during any single event is also a function of rain, at least in part. When rains are minimally adequate, the proportion of tubercules that flower is small; when rains are relatively large, many tub- ercules bloom (Fig. 3). On the other hand, even heavy rains do not induce all potentially fertile tub- ercules to flower during a single event. Because some tubercules are withheld, plants can flower several times a year, given appropriate conditions. Fruit set of M. grahamii and certain other small cacti is relatively high (Table 2). In the case of Echinomastus, fruit set apparently is not affected by pollinator abundance or effectiveness (Johnson 1992). This might not be true for M. grahamii. The results of the caging experiment suggest that plants risk poor fruit set if they happen to flower when pollinator populations are low or when pollinators preferentially visit other species. This does happen at least occasionally, as in 1996 when visitation on one day was essentially nil. Withholding of tuber- cules might allow plants to spread the risk across the entire flowering season, thus increasing the probability that flowering will coincide with polli- nator activity. Thanks to S. P. McLaughlin for reading the manuscript and for providing data from his greenhouse studies. LITERATURE CITED AGUILAR, R. P., T. R. VAN DEVENDER, AND R. S. FELGER. 2000. Cactaceas de Sonora, México: su diversidad, uso y conservacion. Arizona-Sonora Desert Museum Press, Tucson, AZ. ANDERSON, E. FE 2001. The cactus family. Timber Press, Portland, OR. BENNETT, P., R. R. JOHNSON, AND M. R. KUNZMANN. 1986. Cactus collection factors of interest to resource man- agers. Pp. 215-223 in T. S. Elias (ed.), Conservation and management of rare and endangered plants. Cal- ifornia Native Plant Society, Sacramento, CA. Bropy, A. K. 1997. Effects of pollinators, herbivores, and seed predators on flowering phenology. Ecology 78: 1624-1631. GiBsON, A. C. AND P. S. NOBEL. 1986. The cactus primer. Harvard University Press, Cambridge, MA. GOLDBERG, D. E. AND R. M. TURNER. 1986. Vegetation change and plant demography in permanent plots in the Sonoran Desert. Ecology 67:695-—712. GRANT, V. AND W. A. CONNELL. 1979. The association between Carpophilus beetles and cactus flowers. Plant Systematics and Evolution 133:99—102. GRANT, V. AND K. A. GRANT. 1979. The pollination spec- trum in the southwestern American cactus flora. Plant Systematics and Evolution 133:29-—37. JOHNSON, R. A. 1992. Pollination and reproductive ecol- ogy of acuna cactus, Echinomastus erectocentrus vat. acunensis (Cactaceae). International Journal of Plant Science 153:400—408. May, R. W. 1994. The ecology of Sclerocactus polyan- cistrus (Cactaceae) in California and Nevada. Desert Plants 11:6—22. NABHAN, G. P., E. SAUCEDO M., P. OLWELL, P. WARREN, 206 W. Hopcson, C. GALINDO, R. BITTMAN, AND S. AN- DERSON. 1989. Plants at risk in the Sonoran Desert: an international concern. Agave 3:14—15. NewstTroM, L. E., G. W. FRANKIE, H. G. BAKER, AND R. K. COLWELL. 1994. Diversity of long-term flowering patterns. Pp. 142-160 in L. A. McDade (ed.), La Sel- va: ecology and natural history of a neotropical rain forest. University of Chicago Press, Chicago, IL. PRIMACK, R. B. 1985. Patterns of flowering phenology in communities, populations, individuals, and single flowers. Pp. 571-593 in J. White (ed.), The popula- tion structure of vegetation. Dr. W. Junk Publishers, Dordrecht, The Netherlands. MADRONO [Vol. 49 RATHCKE, B. AND E. P. LAcEy. 1985. Phenological patterns of terrestrial plants. Annual Review of Ecology and Systematics 16:179—214. ROLLER, P. S. 1996. Distribution, growth, and reproduction of Pima Pineapple Cactus (Coryphantha scheeri Kuntz var. robustispina Schott). M.S. thesis. Univer- sity of Arizona, Tucson, AZ. SCHMALZEL, R. J., E W. REICHENBACHER, AND S. RUTMAN. 1995. Demographic study of the rare Coryphantha robbinsorum (Cactaceae) in southeastern Arizona. Madrono 42:332—348. SHREVE, FE AND I. L. WiGGINs. 1964. Vegetation and flora of the Sonoran Desert. Stanford University Press, Palo Alto, CA. Maprono, Vol. 49, No. 4, pp. 207-227, 2002 THE EFFECT OF FIRE AND COLD TREATMENTS ON SEED GERMINATION OF ANNUAL AND PERENNIAL POPULATIONS OF ESCHSCHOLZIA CALIFORNICA (PAPAVERACEAE) IN SOUTHERN CALIFORNIA ARLEE M. MOoNnrtTALVO!, LAURA J. FEIST-ALVEY’, AND CATHERINE E. KOEHLER? Department of Botany and Plant Sciences and Center for Conservation Biology, University of California, Riverside, CA 92521 ABSTRACT Throughout its native range, the California poppy, Eschscholzia californica, exhibits substantial mor- phological and life-history variation, including variation in seed dormancy and ability to perennate. Pop- ulations from xeric southern California habitats have high seed dormancy over a range of habitats that span from coast to desert and across vegetation types of varying fire frequency. Understanding variation in the cues that break dormancy in this species is especially important to managing natural populations with prescribed fire, and in production and use of local ecotypes for restoration, erosion control, and ecological landscaping. We explored the influence of sequential treatments of low temperature and com- ponents of fire (heat, dry smoke, smoke water, a commercially concentrated smoke water we call “‘liquid smoke’’, charrate, and nitrate) on seed germination for both annual and perennial populations and com- pared results to those of widely used domesticated seeds. We also examined the effect of light and seed age. Domesticated seeds had no seed dormancy and, except for heat-treated seeds, germination was close to 100% across treatments, including water controls. In contrast, seeds of all wild southern California populations showed some dormancy, germination was highly conditional on test factors, and light inhib- ited germination. We found differences in dormancy rates among wild populations and years since seed collection, with annuals having higher dormancy than perennials in the first year following collection but not after aging > two years. Of the fire treatments, heat (85°C for 10 min), or heat plus smoke, resulted in significantly reduced germination and viability of all populations tested, including domesticated seeds. All smoke treatments significantly improved germination of dormant-seeded populations over water con- trols, but neither nitrate, water soaking, charrate, nor cold treatment alone broke dormancy. In the absence of cold treatment, both liquid and dry smoke yielded higher germination than controls in seeds aged 8— 10 months (65-95% for liquid smoke, 21—60% for dry smoke, vs. 14-59% for controls). Moist cold treatment (3—9°C) by itself did not usually break seed dormancy but it did act synergistically to increase germination of smoke-treated seeds and did not harm controls. In contrast, for most populations colder pretreatment (~2°C) resulted in a small decrease in germination of water controls but not in seeds smoke- treated before cold treatment. Seed age affected germination of controls and the ability of smoke to break dormancy. Germination of controls and smoke-treated seeds increased between 2 and 4 mo of aging in the lab, with no further increase at 8 mo. Dormancy of controls was substantially higher in seeds aged in the lab > 27 months from collection relative to seeds aged 8—10 months (n = 7 and 5 populations, means = 92% and 63% dormant, respectively). Smoke succeeded in breaking dormancy of older seeds to half the extent as in younger seeds, suggesting either a decline in germinability as seeds degrade, induction of a deep secondary dormancy, or both. Given the large differences between domesticated and wild populations in dormancy and germination requirements, and that seed dormancy is probably heritable and adaptive, non-dormant domesticated seeds are not appropriate for restoration, especially in xeric environments that naturally support plants with dormant seeds. Key Words: Eschscholzia, fire, germination treatments, life-history variation, restoration, seed dormancy, smoke, stratification. As the use of native plants for revegetation and landscaping gains popularity, and as restoration projects become more prevalent and species-inclu- sive, understanding seed germination biology be- ' Author for correspondence, e-mail: montalvo @ citrus. ucr.edu * Current address: Remediation Division, Montana De- partment of Environmental Quality, PO. Box 200901, He- lena, MT 59620-0901. * Current address: University of California Davis— McLaughlin Reserve, 26775 Morgan Valley Rd., Lower Lake, CA 95457-9411. comes essential from both basic and applied eco- logical perspectives. In warm and dry climates, na- tive seeds can be difficult to germinate because of dormancy, and they often require very specific ger- mination cues. As is the case with many traits, both dormancy and response to dormancy-breaking sig- nals can be locally adaptive and can differ dramat- ically within species, especially when they exist in a wide variety of habitats (Cruden 1974; Capon et al. 1978; Keeley 1986; Meyer et al. 1990; Meyer and Monson 1992). If cues that break seed dor- mancy are identified, then practitioners can use the information to maximize germination of out-plant- 208 ed seeds and of seeds used for agricultural increase, thus limiting selection against genes conferring dor- mancy. Seed increase encourages use of local seed sources for restoration, landscaping, and roadside erosion control, thereby minimizing mismatching of important adaptive traits to planting location. Mismatches can decrease project success and intro- duce maladapted genes into wild populations (Mon- talvo et al. 1997; Keller et al. 2000; Montalvo and Ellstrand 2000, 2001). The effects of fire on germination response are so common that it is worthwhile to include fire treatments in studies of seed dormancy, especially for species with close relatives known to be fire followers. Seeds of many species in Mediterranean climates, where fire is common, germinate in re- sponse to different fire-related cues (Keeley and Keeley 1987; Keeley 1991; Roche et al. 1997a, b, 1998). Also, as prescribed burning becomes an in- creasingly important management tool, it is impor- tant to examine the effects of fire on seeds, and to determine if populations from different environ- ments have different requirements and vulnerabili- ties. We chose California poppy (Eschscholzia cali- fornica Cham.) as a model species for studying var- iation in seed dormancy and response to different germination cues. California poppy occupies many habitats throughout its native range from Baja Cal- ifornia to southern Washington state, and is native to a diverse range of climates, edaphic environ- ments, and plant communities that vary in fire fre- quency, from coastal sand dunes to inland deserts, including fire-prone coastal sage scrub, chaparral, and grasslands. Populations exhibit a wide spec- trum of morphologies and life histories, varying in flower color, size, seed dormancy, and other attri- butes, and can be annual (semelparous), long-lived perennial (iteroparous), or a facultative annual (Cook 1962). In addition, this species has promi- nent economic importance as the California state flower and is frequently used in restoration and re- vegetation. Information about variation in its ger- mination response to the effects of fire and other factors is essential for long-term success of projects that utilize prescribed burning for restoration, for commercial seed production, and for conservation practice. In southern California, native poppies (both an- nuals and perennials) germinate during the rainy season in mid winter; plants flower in late winter to spring, and produce seeds in the spring (ate April to early June), although some perennials can defer flowering to the second spring and produce seeds over a longer season. Seeds appear to remain dormant (or conditionally dormant) until the next winter rains, which prevents early germination after infrequent summer storms and subsequent death from desiccation in the hot, dry summer environ- ment. Thus plants appear to fit the ““winter annual and perennial’”’ syndrome (Baskin and Baskin 1998, MADRONO [Vol. 49 p. 54-56), but it is unknown whether the seeds fol- low an annual cycle of summer dormancy followed by non-dormancy or conditional dormancy during the winter. Based on the small linear embryo of California poppy seeds, seeds have been assumed to have morphophysiological dormancy in which embryos complete their maturation after seed dis- persal and an environmental cue is required to break physiological dormancy (Martin 1946; Bas- kin and Baskin 1998). It is unknown if embryo growth and dormancy break require the same or different conditions. In interior shrub- and grassland habitats of south- ern California, prolific displays of poppies in the absence of fire are periodic and tend to be separated by many years. In some years, flowering of annual populations is absent or scant and can be followed by boom years, demonstrating the existence of a substantial seed bank. This suggests that seeds may cycle through dormancy and conditional dormancy depending on environmental conditions. In addi- tion, the conspicuous presence of poppies on recent burns when blooming is poor elsewhere (A. Mon- talvo personal observation, G. Hund, R. Noll, J. Crossman personal communication) suggests that, at least in these habitats, seeds survive fire and some component of fire aids in breaking seed dor- mancy. In California, fire and fire components have been shown to break seed dormancy in many “‘fire-fol- lowing” species, especially in sage scrub and chap- arral communities (Keeley 1991). For example, seed dormancy of Emmenanthe penduliflora Benth., Sal- via columbariae Benth., S. mellifera Greene, Pha- celia grandiflora (Benth.) Gray, P. minor (Harv.) Thell., and Lotus scoparius (Nutt.) Ottley is broken by heat, smoke, or other components of fire (Keeley 1991; Keeley and Fotheringham 1998a, b). Each of these species occurs in a number of different plant communities, frequently concomitant with California poppy (Munz and Keck 1968). In studies of intra- specific germination response to heat, very different patterns were found among populations of perennial S. mellifera (Keeley 1986) and annual S. columbar- tae (Capon et al. 1978) from desert habitats vs. chap- arral and coastal sage scrub. Given that the distri- bution of California poppy is even more diverse than for these Salvia species, its germination response to fire is likely at least as complex. The response of poppy seed germination to components of fire has not been studied, although prescribed burning is be- ing used to manage portions of its native habitat, including the Antelope Valley California Poppy Re- serve (AVCPR, www.calparksmojave.com/poppy; www.parks.ca.gov/parkindex, J. Crossman, personal communication). Data on the general germination biology of Cal- ifornia poppy are scant, and few authors have iden- tified use of wild rather than domestic seeds in their work (e.g., Cook 1961, 1962; Fox et al. 1995). Cook (1961, 1962) documented geographic varia- 2002] TABLE 1. MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 209 SEED POPULATIONS USED IN SEED GERMINATION EXPERIMENTS. Abbreviations: Pop = population; A = annual; P = perennial. Source, locality, and life-history strategy are listed for each study population. Source of seeds and collection date S&S Seeds (Carpinteria, CA) Lot #T8979, ~May 1998 L. Feist, May 25, 2000 Stover Seed Co. (Los Angeles, CA), harvested for 2000 market J. Crossman, May 1995 G. Fox, ~May 1996 L. Feist, May 17, 2000 A. Montalvo and L. Feist, May 27, 2001 S&S Seeds (Carpinteria, CA), Lot #87383, ~May 1997 R. Noll, S&S Seeds (Carpinteria, CA), Lot #T1008, Apr. 29—May 11, 1998 L. Feist, May 11—23, 2000 A. Montalvo, L. Feist, May 8-13, 2001 G. Fox, ~May 1996 L. Feist, May 25 and June 6, 2000 A. Montalvo and L. Feist, May 27, 2001 R. Noll, S&S Seeds (Carpinteria, CA), Lot #T1022, May 7-9, 1998 R. Noll, S&S Seeds (Carpinteria, CA), Lot #V 1022, May 8-11, 2000 A. Montalvo and J. Skillman, May 19, 2001 Pop Life Code history Collection locality LC98 P Antelope Valley area, Lancaster, CA AVO0O Pp Antelope Valley area 2, CA Dom P Cultivar domesticated in Salem, OR 25-30 generations (origin s. California) PR95 A California State Parks, Antelope Valley Cali- fornia Poppy Reserve, CA (AVCPR) PR96 A as in PR95 PROO A as in PR95 PROI A as in PR95 CB97 A Carlsbad, CA NH98 A North Hills of Western Riverside County Multi-Species Reserve, CA NHO0OO A as in NH98 NHO1 A as in NH98 FM96 E Fairmont Butte (s. of AVCPR) FMO0O Pp as in FM96 FMO1 Pp as in FM96 EM98 A Vicinity of Estelle Mt., ne Lake Elsinore, Riverside Co,. CA EMO00 A as in EM98 SR101 lf Serpentine site, Sedgwick Ranch Reserve, Santa Barbara Co., CA RVO1 P Riverside, Riverside Co., CA, domesticated population tion in seed dormancy that was loosely correlated with habitat and longevity; annual and perennial populations from xeric sites had mostly dormant seeds, whereas perennial populations from mesic sites had little to no dormancy. He found that dor- mancy could sometimes be broken with gibberellic acid (GA3), but he did not identify the natural cues that break dormancy. Here we assessed the germination response of E. californica seeds from several southern California populations to various components of fire and other potential dormancy breaking conditions. We includ- ed seeds from both annual and perennial wild pop- ulations, neighboring and distant populations, and a variety of climate regions including coastal, in- land, and desert. We examined whether the popu- lations varied in seed dormancy, effect of seed ag- ing, and in germination response to a variety of fire treatments, including heat, smoke, charrate, and ni- trogen. Poppies do not usually germinate until well into the cold season (December—February). Be- cause many GA3 responsive seeds also respond to cold treatment, we also explored the effect on ger- mination of exposure to a short period of cold tem- peratures (e.g., Roche et al. 1998), using varying temperatures, length of exposure to cold, and mois- ture levels during exposure to cold. A. Montalvo May 27, 2001 METHODS Study populations. All wild populations were from southern California and represented both an- nual and perennial life-histories (Table 1). Seed col- lection codes designate site of collection followed by year of collection (1.e., 97 for 1997; 00 for 2000 and so on). Seeds were collected usually in May just before capsules exploded. Habitats of wild source populations varied as to vegetation type, precipitation, edaphic features, and other factors. Four study sites (LC, PR, FM, and AV) were grass- land mixed with forbs in the Antelope Valley of the upper Mojave Desert at or near AVCPR, and two sites (SR) were shrub-grassland ecotones in the foothills of Santa Barbara Co. Of these, only site PR (from AVCPR) supported annuals. The remain- ing annual populations were from sage scrub hab- itat in Riverside (NH, EM), and coastal San Diego Counties (CB). Mean temperatures of winter months vary among sites (Table 2), and were used to guide choice of chilling and incubation temper- atures. The domesticated population (Dom) has been repeatedly planted and harvested for com- mercial seed production in Oregon for at least 20 years. The original source population for Dom is unknown, but thought to have been from southern California. The garden population RVO1 was plant- 210 MADRONO [Vol. 49 TABLE 2. MEAN DarILy MINIMUM, MAXIMUM, AND MEDIAN TEMPERATURES FOR SOUTHERN CALIFORNIA WEATHER STA- TIONS NEAR STUDY POPULATIONS. Closest weather Study station location population Lake Cachuma (1952-2000) SR Lancaster (1945-1960) AV, FM, LC, PR San Diego (1940-1960) CB San Jacinto (1948—2000) NH Sun City (1973-2000) EM ed before 1990 from an unknown commercial seed source. Seed germination experiments. Six experiments were conducted, with each building on the infor- mation gained from previous experiments. General procedures are described below. Details of popu- lations and experiments are in Table 3, including seed age at time of incubation. General protocol. After we collected or re- ceived seeds, we stored them in the lab at ~22 + 3°C. Before we obtained seeds from other sources, they had been stored under variable conditions: S&S Seeds stored seeds at ambient conditions in warehouse in coastal, Santa Barbara Co.; Fox and Crossman stored seeds in offices; domesticated seeds were stored under unknown conditions before purchase in June 2000. All wild site collections, including those from S&S Seeds, were document- ed. For all experiments, we chose visibly undam- aged, plump seeds of uniform shape, and which had obtained a brown to blackish brown color. Seeds were always randomly assigned to treatments (de- scribed below). For incubation, one filter paper (Whatman No. 1, 90 mm) was used per petri-dish (Fisher, polystyrene, 90 X 15 mm). Immediately before incubation, for treatments Con, S, H, SH, and Q (below), we placed seeds on filter paper moistened with 2 ml of pH7.5 filter sterilized tap water (hereafter ““water’’). For treatments L, W, and N, we placed seeds on filter paper moistened with 2 ml of the treatment solution. Unless otherwise indicated, incubation then occurred under a diurnal cycle of 12 hr/22°C and 12 hr/12°C (henceforth 22/ 12°C). We incubated seeds in winter (excepting Exp. | & 2) when they would be germinating in nature to control for any seasonal cycling of dor- mancy under shelf conditions (see Baskin and Bas- kin 1998). We aged seeds at least 8 mo in the lab except when testing for after-ripening (Exp. 6), considered here as a period of embryo maturation following seed harvest that occurs before seeds can germinate (Nikolaeva 1969). For cold treatment, seeds were placed in dishes between two filter papers and either left dry or moistened (see below). Before cold treatment or in- cubation, dishes were individually wrapped in alu- Mean min/max temperature °C (median temperature) December 3.6/19.3 (11.5) —1.8/14.9 (6.5) 9.5/19.6 (14.5) 1.1/19.4 (10.3) 1.4/19.8 (10.6) January 3.7/18.5 (11.1) ~1.9/13.8 (5.9) 8.1/18.3 (13.2) 1.1/18.4 (9.7) 2.4/18.9 (10.7) minum foil to exclude light and placed in sealed plastic bags to prevent moisture loss or gas ex- change between treatments. Cold treatments and in- cubation took place in the dark because light was reported to inhibit germination of California poppy (Goldthwaite et al. 1971). We confirmed that light inhibits germination in wild-collected seeds but not in domesticated seeds. We tested the effect of light on germination by in- cubating 100 moistened seeds from each of four wild and one domesticated population in light (no foil wrap) vs. dark (foil wrapped) under alternating 11 hr days and 13 hr nights (temperature ranged 10—20°C). After 19 days, mean germination under light was 11.7%, 3.6%, 3.2%, 5.7%, and 99%, while germination in the dark was 29.3%, 16.5%, 25.8%, 100%, and 100% for populations EMOO, PROI, PalaOl, RMO1, and Dom, respectively (RMO1 = perennial from coastal San Luis Obispo County; PalaOl = annual from inland San Diego County). At harvest, we scored radicle emergence of at least | mm as successful germination. Any remain- ing, ungerminated seeds were counted and assayed for viability with 2,3,5-triphenyl tetrazolium chlo- ride (TTC) (Fischer), a metabolic indicator dye. Seeds were pierced with a probe, stained by soak- ing overnight in a 1.0% TTC solution (Kitchen and Meyer 1992), then dissected under a dissecting mi- croscope. A seed was scored as viable if more than two thirds of its embryo stained dark pink or red. Live, ungerminated seeds were considered dor- mant. The proportion of viable seeds in each dish was ((#germinated seeds + #viable ungerminated seeds)/total # seeds) and proportion germinated was (#germinated seeds/#viable seeds), thereby stan- dardizing germination to the response of viable seeds. Fire treatments. Dry fire treatments (S, H, SH) were given before any temperature treatments whereas wet fire treatments (N, L, W, Q) were giv- en after temperature treatments in the first two ex- periments. Thereafter, wet fire treatments were giv- en before temperature treatments. S (dry smoke): Smoke stimulates germination in many species and can be applied by various meth- ods. 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We placed dry seeds in small plastic dishes in a large glass chamber. A 500 ml airtight metal can was filled 2/3 full with equal portions of air-dried Avena fatua L., Bromus madritensis L., B. diandrus Roth, Lessingia filaginifolia (Hook. & Arn.) M. A. Lane, and Nassella pulchra (A. Hitchc.) Barkworth. The vegetation was ignited, and the smoke was transferred into the glass chamber with forced air through a long tube. The chamber was filled with the cooled smoke for 30 seconds, then sealed, and seeds were smoked for five minutes. Our prelimi- nary experiments with domesticated poppy seeds showed that smoking up to 15 minutes does not affect seed viability. H (heat): A brief heat shock has been shown to enhance germination of numerous species (Keeley 1991; Baskin and Baskin 1998), while killing or inhibiting germination of heat-sensitive seeds. To explore E. californica’s response to temperatures similar to those that may be experienced by buried seeds during a fire, we heated dry seeds in open Pyrex beakers in a forced air oven at 85°C for 10 minutes. Preliminary trials using a temperature as- say range of 70—115°C demonstrated that viability of both wild and domesticated seeds decreased at 115°C, but not all seed populations lost viability at 85°C/10 min (Feist and Montalvo, unpublished data). SH (smoke + heat): A random subset of dry- smoked seeds was heated as in H. N (nitrogen): Levels of available N as nitrate have been shown to increase after fire (Sweeney 1956; Franco-Vizcaino and Sosa-Ramirez 1997), a chemical shift which enhances germination in some species (Thanos and Rundel 1995; Baskin and Bas- kin 1998). Also, E. californica tends to respond positively to disturbance, which in turn is positively correlated with nitrogen availability. We used a so- lution of 10 mM KNO, (Fischer) which has been used successfully to stimulate germination in some fire-following species (Thanos and Rundel 1995; Keeley and Fotheringham, 1998b) to moisten seeds. L (leached charrate): A leachate of the charred remains of plant material enhances germination in some species (Keeley 1991), and simulates water transporting charred plant particles to the seed. We prepared charrate for adding to seeds by burning equal portions of air-dried vegetation (same as in S) until blackened but not ashed. This material was ground and added to water (5 g per 100 ml), stirred overnight, then filtered through several layers of cheesecloth (modified from Keeley 1991). W (smoke water): In this smoke treatment, wa- ter picks up smoke particles/chemicals and transfers them to the seed (Keeley and Fotheringham 1998b). We filled an airtight 500 ml can with one of the vegetation types (see S above), set the material on fire, and forced the smoke to bubble though 1 liter of water via a tube. Dried material of each species MADRONO [Vol. 49 (S above) was burned in series until all the material was completely blackened (1—5 minutes). Q (liquid smoke): Commercially produced “‘liq- uid smoke’”’ products enhance germination in many plant species, and one such product has been used successfully in Australia to increase germination on mine reclamation sites (Roche et al. 1997b). We obtained commercial liquid smoke (Regen 2000 Smokemaster® Seed Germination Solution, Regen, Glasgow, KY) made from passing smoke through water and concentrating it. In Exp. 1-2, we soaked seeds after their cold treatments (or after dry control) in a dilution of 1: 50 (Regen 2000:water) for 20 hours. This treatment was modified for subsequent experiments. In the first modification (mQ-1), seeds were soaked in a dilution of 1:50 for 12 hours, then air dried at room temperature. Seeds were then subjected to appro- priate cold treatments after smoke treatment (Exp. 3—4). The second modification (mQ-2) explored the effects of liquid smoke concentration (Exp. 5, be- low). Although the manufacturer recommends us- ing a 1:10 dilution, a 1:50 dilution had been used in order to decrease the risk of harming seeds with high liquid smoke concentrations. To assess the ef- fects of dilution strength on germination, we soaked seeds 12 hr in either a 1:50, 1:25. or 1:10 dilution, then transferred wet seeds to the appropriate cold- shock treatment (see below). Con (control for fire treatments): No fire treat- ments were given, and seeds were subjected to the appropriate temperature and moisture treatments (below). Temperature and moisture treatments. Cold: Seeds in petri dishes between two filter pa- pers were subjected to prolonged cold temperatures in a cold chamber (ranging from two to eight weeks) while either moist or dry. Seeds were moist- ened with 2 ml of fluid (moist-cold), 3 ml of fluid (wet-cold), or left dry (dry-cold). Wet-cold treat- ment provided free water for rapid imbibing, moist- cold provided for slower imbibing and high relative humidity, while the dry method tested whether pop- py seeds can be affected by cold treatment when dry. In addition, Exp. 5 tested the effects of a brief cold-shock (48 hours at 2°C) instead of prolonged cold treatment. Cold treatment was intended to occur near or be- low 4°C. Problems with one cold room resulted in higher cold-treatment temperatures during Exp. 3 (see below), averaging near 7°C (range 4—9°C), which resulted in a positive germination response and provided the impetus to test variation in incu- bation temperature in Exp. 5. No-cold (control for cold and moisture treat- ments): For dry, no-cold conditions, henceforth ‘“‘no-cold’’, we stored seeds dry at room tempera- ture until moistened and incubated. Experiments. All experiments included a control treatment, for which seeds received no fire or cold 2002] treatments prior to incubation. At the time of in- cubation, those seeds collected in spring 2001 were between 2—7.5 months old (henceforth ‘“‘fresh seeds’’), seeds collected in 2000 were between 8— 10 months old (henceforth “‘young seed’’), and all other wild seeds were > two years old (henceforth ‘“‘aged seed’’). Experiment | and Exp. 2 were run concurrently and were overlapping subsets of one experiment. In Exp. 1, we examined germination response of three ‘“‘aged seed”’ populations and domesticated seeds to cold treatment at three different moisture levels (wet, moist, and dry) in combination with five ‘“‘wet’’ fire-effects treatments (Table 3). The wet fire treatments were applied after seeds had been cold- treated. In addition, for Exp. 2 we added three “‘dry”’ fire treatments (S, H, and SH) under three cold treat- ments (moist-cold, dry-cold, no-cold treatment) and on seeds of two additional wild populations (PR95 and FM96, for a total of six populations). The ap- plication of dry fire treatments before seeds were chilled, simulated the natural sequence of events in the wild. The chilling temperature ranged from 2-— 4°C; most often close to 2°. For both Exp. 1 and Exp. 2, incubating seeds were checked using dim green lights after 3, 8, 13, and 20 d of incubation. No increase in germination was seen after 13 days. In Exp. 3, we investigated five aged and one young seed population, increased the number of replicates from three to five petri-dishes to increase statistical power, and examined the response of wet and dry smoke (mQ-1 and S) to a longer 8 wk cold treatment and a more natural sequence of smoke and cold events (Table 3). Both the S and mQ treat- ments were applied before seeds were subjected to moist cold treatments. The cold room cycled be- tween 4—9°C, and was mostly just below 7°C. After cold treatment, any germination was tallied, and ungerminated seeds were transferred into new dish- es with fresh filter paper, moistened with 2 ml wa- ter, and incubated for 20 days before scoring ger- mination and viability. Experiment 4 assessed germination of young seeds collected in the same year to investigate their response to smoke and cold treatments (Table 3). Data from Exp. 3 suggested that young seeds (<1 yr) may be less dormant or have dormancy more easily broken than aged seeds (2+ yr) which could complicate experiments with mixed age seeds. Ex- tended shelf storage can trigger secondary dorman- cy in some species while decreasing dormancy in others (Roche et al. 1997a; Baskin and Baskin 1998). In Exp. 5 we tested different concentrations of liquid smoke (mQ-2), different incubation temper- atures, and the effect of a brief cold-shock on ger- mination and viability of two populations, one with young seeds and one with aged seeds (Table 3). Different incubation temperatures (22/12, 18/6, and 7.5°C constant) were tested to determine if lower MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 2B, incubation temperatures improved germination, given that germination was possible and sometimes higher at quite low temperatures (Exp. 3). In ad- dition, we subjected half the seeds to a 48 hr cold- shock at 2°C instead of prolonged cold because short cold treatments are sometimes sufficient for either breaking or inducing dormancy and are eco- logically realistic within E. californica’s range. Fi- nally, to discern between the possible effects of soaking which may leach germination inhibitory chemicals from seeds, and the effects of wet fire treatments, we added a wet-control treatment (12 hr soak in water). We used annual population EMOO for the full Exp. 5. In addition, seeds from the deeply dormant NH98 population were run with only a wet-control and with 18/6°C as the incubation temperature (based on incubation temperature with highest ger- mination in Exp. 4) to see if germination could be improved with higher or lower concentration of lig- uid smoke. We ran Exp. 6 to test if recently collected “fresh”? wild and domesticated California poppy seeds germinate at different rates with and without smoke treatment over the course of several months after fruit dehiscence. If seeds require lengthy after- ripening or if in the lab they experience an annual cycle of dormancy followed by conditional dor- mancy or loss of dormancy as expected under nat- ural conditions, then these behaviors could affect the outcome of germination experiments run at dif- ferent seed ages or times of the year. We collected seeds into paper envelopes in May 2001 from four wild populations (perennial FM and SR; annual NH and PR, Table 3), and from garden plants in Riv- erside (RV—started from commercial seed source > 10 years earlier, Table 3). Six weeks after col- lection and shelf storage, we randomly sorted seeds into microcentrifuge tubes, and then tested at 2, 4, and 7.5 months (herein “‘8’’ mo) following seed harvest. We incubated controls and liquid smoke- treated seeds (mQ-1) in the dark at alternating 18°C/6°C for 11/13 hours for 15 days and scored germination and viability. Subsamples of smoked seeds aged 4 mo and 8 mo were given moist-cold treatments (4 wk at 3.5°C and 2 wk at 3°C, respec- tively) before incubation. Data analysis. Data were analyzed with ANOVA using Proc GLM of SAS (Release 6.12). In all mod- els, source POPULATION was a random effect while COLD (e.g., cold vs no-cold), FIRE (e.g., smoke, heat, nitrate, water, etc.), or AGE treatments were fixed effects. Response variables included proportion of viable seeds germinated/dish, and the proportion of viable seeds out of total seeds/dish. Dishes were the replicates. Before analysis, all data were angularly transformed (arcsin(proportion)'”) to enhance the normality of the residuals. We used either Duncan’s Multiple Range test or Tukey’s test for posteriori comparisons among means. In mod- 214 els where we found significant interactions among main effects, we ran separate analyses on each pop- ulation or treatment depending on structure of the interaction and the particular question. Given the mixed model ANOVAs, we used the RANDOM statement in SAS to calculate the denominator mean squares for F-tests using the Satterwaite mod- el. RESULTS General patterns. Over experiments, seed dor- mancy (Table 4) and the effect of potential dor- mancy breaking treatments varied substantially among years of collection and among populations (Tables 5, 6; Fig. 1-5). The domesticated commer- cial (Dom) and garden (RV) populations were the only ones with no seed dormancy (Table 4). Other populations ranged from 41% to 100% dormant un- der control conditions. In general, young seeds col- lected in 2000 had higher germination in controls and higher germination following treatment with liquid smoke than aged seeds (mean 65% vs. 92%; Table 6). There were differences in seed dormancy among young (year 2000) collections from the Antelope Valley, with much lower dormancy of the perennial AVOO0O and FMOO than the annual PROO. Aged seeds had uniformly high dormancy (range 93—100%). Seed viability also varied among wild collections and ranged from 54—100% following treatments (Table 4). Populations NH98 and FM96 showed different levels of viability between experiments, possibly because of differences among technicians in sorting bulk seeds to be used in experiments. The range in viability shows the importance of basing % germination on live seeds rather than total seeds. There were highly significant effects of fire and cold treatments and significant population effects. In addition, the many significant two and three way interactions between FIRE, COLD, AGE, and POP- ULATION main effects (Table 5) indicate variation among populations in response to at least some treatments. Because of this complexity, we present results of each experiment separately and break up analyses to examine interactions and effects of fire, cold, and age treatments. Given their lack of seed dormancy, we did not statistically compare Dom and RV with wild pop- ulations. The domesticated seeds had nearly 100% germination and viability in all treatments except those involving heat. In addition, unlike the dor- mant-seeded populations, Dom seeds germinated during cold treatment at 2—4°C. Separate analysis of population Dom seeds under the no-cold regime showed that there was no significant difference among the Con, S, N, W, L, or Q treatments. How- ever the H and SH treatments reduced germination significantly suggesting that heat treatment inhibit- ed germination and even killed some seeds. Via- bility of seeds from the H and SH treatments ranged from 64—100%, compared to 100% in controls. MADRONO [Vol. 49 Experiment 1—Effect of wet-fire and cold treat- ments (moist-cold, wet-cold, dry-cold, and no-cold). Analysis of germination from the three study pop- ulations of different regions (annual NH98, peren- nial LC98, and annual CB97) revealed significant main effects and interactions except POPULATION and FIRE x POPULATION (Table 5). When ger- mination was analyzed for each population sepa- rately, FIRE, COLD and FIRE X COLD were still significant for each population (all P values = 0.0113). This significant interaction shows that each population responded to the set of treatments differently, obscuring whether any particular cold treatment or fire treatment resulted in the highest germination. However, for all populations, the no- cold regime resulted in the highest mean germina- tion (Fig. 1), suggesting that the 2°C cold treatment was cold enough to slightly inhibit germination in the three populations examined. We also analyzed germination response separate- ly by level of cold treatment and found no signifi- cant FIRE < POPULATION interactions, and in the wet-cold analysis we found no significant FIRE or POPULATION effects (Fig. 1). In the moist-cold treatment, annual NH98 had significantly higher germination than perennial LC98 and annual CB97 seed populations, while in the no-cold regime CB97 seeds had significantly higher germination than LC98 and NH98 (Fig. 1). Inland NH98 seeds ap- pear to germinate under cold conditions more read- ily than coastal CB97 seeds. There were significant effects of FIRE treatment only in the dry-cold and no-cold regimes, potentially because liquid fire treatments were applied after chilling in moist-cold and wet-cold. In no-cold and dry-cold, treatment Q resulted in the highest germination, although Q and W were not statistically different under the no-cold control conditions (Fig. 1). With ANOVA of Experiment 1, we examined the effect of treatments on seed viability to reveal if some treatments harmed seeds. We found no sig- nificant main effects, but the COLD X FIRE xX POPULATION interaction was significant (F534 129 = 1.67; P < 0.04), suggesting that seed viability of different populations was affected differently by the cold and fire treatments. After running viability data separately by population, no significant effects of COLD or FIRE treatments were found for CB97. Population LC98 showed a significant effect of FIRE treatment (Fy 3, = 3.07; P = 0.027) and a FIRE X COLD interaction (Fy, 4 = 2.26; P = 0.028). COLD had a significant effect on NH98 (F’; 49 = 6.76; P < 0.001), with the no-cold regime more viable than all other cold treatments. No other effects were significant. Overall, the wet-cold treat- ment resulted in the lowest germination in NH98 (Sigal): When viability was analyzed for each cold re- gime separately, the only significant effect of POP- ULATION was in the no-cold treatment (Fy , = 4.73; P = 0.044), with NH98 having greater via- MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 25 2002] (L'7) LO (L'S) €6 (€'L) 96 ($9) 76 (L'T) 86 (10'0) SL (T°0) 08 (Sl) 86 (77) $6 (91) 78 (SO) 16 (9°8) PL (Ey) $6 (L'9) 7S (Ey) L8 (6°S) CL (69) TL (0) OO! 7 a) II ae (9°01) 8S = (TI 1) 6S (6'€) 68 = WD) Bt = ra (v'8) 76 (€°7) $8 as, aie (8°T1) LS (v'8) 98 ai (OCI) €9 ag (9°E€1) 76 (9°71) 89 (8°0) 9r (L'0) Iv (€°S) OL (6'0) 98 (Sr) 76 (O'LT) 87 (SL) 88 (0) OO1 (L’€) 16 (€'b) 66 (L'1) 96 (€°€T) €6 (PST) 88 (L'T1) 76 (6°71) S8 meg << 4.13: P < 0.009), with S producing significantly higher germination. For NH98, mean germination was highest for S, but non-significant (Fig. 2). A Ken- dall’s analysis of the rankings of fire treatment within each population showed a strong association between fire treatment and germination (0.01 < P Mean % germination No-cold Dry Liquid Dry smoke Control smoke smoke 55.4 MB) 37.8 29.4 70.6 52.6 83.3 60.2 68.7 58.9 82.9 49.7 aS 6.3 46.5 44.0 0) 0 3.6 0 48.7 ihe s7/ 65.5 215 24.1 11.9 41.9 19.3 25.6 11.8 33.8 2S = Fh GEN 34.0* — 132 29.9 94.9 oy es 24.8 3.8 10.5 10.7 3.6 4.8 25.6 6.6 735 53.9 69.7 59.4 36.6 9.3 65.2 [5:2 — Sil 94.6 = < 0.001). Treatment S had the highest overall rank- ing, HS and H had the lowest overall rankings (in- dicating heat reduces germination), while all other treatment rankings were very similar. Under the no-cold regime, germination of all populations except FM96 was significantly affected by FIRE treatment (all F,,, => 6.2;.P.= 0.001)..A Kendall’s analysis of the rankings of treatments within each population showed a strong association (P < 0.001) between fire treatment and germina- tion. For all populations, Q produced the highest mean germination, while S and W tied for second place (Fig. 2). Averaged over the five populations, the no-cold controls germinated only 25% as well as seeds treated with liquid smoke (Q). The H and HS treatments were ranked lowest overall. Again, the heat treatments reduced germination. Germi- nation of H treated seeds for the five populations averaged 38% of control seed germination. Q per- formed best in the no-cold regime, while S_ pro- duced the highest germination in the cold regime, again suggesting the effect of Q would improve if applied before chilling. This hypothesis was veri- fied in Exp. 3, below. ANOVA of Exp. 2 viability data showed that POPULATION, FIRE, and COLD were all signifi- cant effects (POP F, ,; = 5.76; P = 0.049; FIRE Fa 93, = 4.53; P = 0.002; COLD F, , = 12.4; P = 0.024). There were no significant interactions (all P >0.11). In the full model, viability was unaffect- ed by all treatments except H and SH, which sig- nificantly decreased viability (Duncan’s Multiple 218 MADRONO [Vol. 49 = 100 Moist-cold o 80 £ 60 E 40 0) © 20 xs (0 Con L N Q WwW Con L N Q WwW Con L N Q W LC98 LC98 LC98 LC98 LC98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 No-cold a c no) —s © = = = ® O x Con L N Q WwW Con L N Q Ww Con L N Q WwW. LC98 LC98 LC98 LC98 LC98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 Dry-cold S 100 = 80 £ 60 E 40 0) O20 0) Con L N Q WwW Con L N Q WwW Con L N Q WwW LC98 LC98 LC98 LC98 AV98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 c 100 Wet-cold ne) — O = = _ ) O xs Con L N Q Ww Con LC98 LC98 LC98 LC98 LC98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 Fic. 1. N Q WwW Con L N Q Ww Experiment 1 backtransformed germination data, analyzed for each population within each cold treatment. Lower and upper case letters represent significant differences between treatments and populations, respectively, based on Tukey’s tests (alpha = 0.05). Within each cold treatment, populations or treatments that share a letter are not significantly different. In moist-cold, NH98 had higher germination than CB97 and LC98, but there were no significant effects of fire treatment. Error bars = 1 SD (n = 3). Abbreviations: Con = control, L = leached charrate, N = nitrogen (KNO,;), Q = liquid smoke, and W = smoke water. Range Test, using alpha = 0.05). Viability of SH and H treated seeds for the five populations aver- aged 6% lower than viability of control seed, com- pounding the effect of reduced germination. Over- all, FM96 had significantly higher viability than all other populations, and CB97 had significantly low- er viability than all but population LC98 (Tukey’s test, alpha = 0.01) (Table 4). For remaining experiments, analysis of seed vi- ability will not be presented in detail. Additional treatments had little affect on seed viability. Experiment 3: Effect of dry smoke, liquid smoke and moist-cold, dry-cold, and no-cold (smoke be- fore cold). In Exp. 3, many seeds germinated un- expectedly during moist cold treatment, including some FIRE treatment controls. For populations that germinated during cold treatment, very few of the remaining seeds germinated after shifting to the warmer incubation period. Therefore, in ANOVA of total % germination, the main effect of COLD represents three treatment levels: moist-cold = in- cubation at 4—9°C; no-cold = incubation at 22/ 12°C; and dry-cold = incubation at 22/12°C follow- ing 8 wk dry cold treatment. ANOVA of total % germination showed a sig- nificant COLD X FIRE X POPULATION interac- tion (Table 5). When data were broken up by pop- ulation, treatment mQ (modified liquid smoke) pro- duced significantly greater germination than the other treatments in all populations (averaging over fire treatments shown in Fig. 3), but there was a significant COLD X FIRE interaction within EM98 and NH98. EM98 and NH98 were the only popu- lations significantly affected by COLD (P < 0.0024), and in both cases moist-cold produced higher germination than dry-cold and no-cold (Fig. 2002] MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 219 = = 100 - MH no-cold (11 moist-cold SR Gy tha ce Bsn a rh ® O 3S ee) 3 = Gay on) tie ste - anQ ow —N ACon4AS,. 1H: SH, L.- Q .W «N ‘= S& 100, m no-cold L] moist-cold = bey. 6 4G be jaa «ab bey BeTA » B B B BE B B £ 80 E ay O 3S To) yop) a Con §$ Hen SH Lk Oni We) | NY jCon-,-'S H -Siy ik O° NV" een S @ no-cold CL} moist-cold = 100 abc abc c_ abc abc ab. . ber 7B A Bi) Bye IB B.» Bie B £ = ® O se ~—m op) jaa) O S 100 , —@ no-cold C1) moist-cold £& g91 ab ab ab ab ab avcab . by JAy. A.B ..AB,. AB..AB.. A: -AB E ® O se ee) ror) a Zz = 12, i) = = ® O 3S ie) D z Gono? Ahrshnl? “GW NecGon «Sy He SH ££ »Q. (Ween Fic. 2. Experiment 2 backtransformed germination data, analyzed separately for moist-cold and no-cold treatments within each population. Lower and upper case letters indicate significant differences (Tukey’s test, alpha = 0.01) occurring between treatments within no-cold and moist-cold regimes, respectively. FIRE treatments that share a letter are not significantly different. Error bars = 1 SD (n = 3). Abbreviations: as in Fig. 1 plus S = dry smoke, H = heat, and SH = smoke plus heat. Evidently, germination decreased as seed age in- creased. 3). In a separate analysis by cold regime, there was a significant interaction of FIRE x POPULATION in no-cold and dry-cold, likely due to populations CB97 and EM98, for example, having much larger differences between mQ and control treatments than population AVOO. However, in all cold treat- ments, mQ produced the best germination, and was highest for AVOO, then EM98, and iowest for PR96. Experiment 4: Effect of dry and liquid smoke vs. cold treatments on ‘‘young”’ seeds. ANOVA of ger- mination data showed that all effects were signifi- cant except COLD X POPULATION and COLD xX TREATMENT X POPULATION (Table 5). Even 220 MADRONO [Vol. 49 8 © 100 = E 60 8 8 40 xe sx 20 ~ E b s i 40 x = © 20 = S e 2 mo before they were capable of dormancy brake and germination (Fig. 6). We also looked at effects of mQ-1 and a cold treatment at 3—3.5°C on 4 and 8 mo-old seeds (Fig. 6). A separate ANOVA of treat- ment (cold + mQ-1 vs. mQ-1) and age (4 vs. 8 mo) on germination, showed no significant effect of cold treatment or age in PROI or FMO1. However, cold significantly increased germination in NHO1 and SRIO1 seeds and decreased germination in SR601. Germination was significantly higher for 4 mo-old seeds in these later three populations. Thus cold treatment had an inconsistent effect on ger- mination relative to just smoke-treated seeds both among age groups and among populations. DISCUSSION Our study populations came from a set of con- trasting southern California environments. We used seeds of both annual and perennial life-histories and with different levels of seed dormancy. We ex- amined whether cold treatment, some specific com- ponents of fire, or some combination enhances ger- mination of dormant seeds, and found important di- rect effects of smoke. This is important because his- torically, fire has been naturally occurring and periodic in both shrub and grasslands of California, (Parsons 1981; Keeley 1991) where poppies occur. Additionally, for thousands of years prior to Euro- pean settlement, indigenous tribes of southern Cal- ifornia managed many areas with fire to encourage growth of animal forage as well as certain food and textile plants (Bean and Lawton 1973; Timbrook et al. 1982; Lewis 1993). With such repeated exposure to fire, we expect that many species that have not been considered fire followers may, nonetheless, re- spond to chemical constituents of fire. Even in de- sert areas that do not have a history of frequent wildfire, the incidence of fire is increasing due to fuel loading by invasive grasses coupled with in- creasing human activity (Brooks 1999), underscor- DDD MADRONO EMO0 @ 22/12 C A A A o> o> A b a a a 100 B B 80 B 5 60 4 2 a b b % Germination pes [o) Cn-D Cn-W 1:50 1:25 1:10 Cn-D Cn-W 1:50 1:25 1:10 NCS NCS NCS NCS NCS CS CS CS CS _ CS EMO00 @ 18/46 C A A A A A A a a a b b b 100 B B 80 B B a a % Germination £& OD oo lon oOo Cn-D Cn-W 1:50 1:25 1:10 Cn-D Cn-W 1:50 1:25 1:10 NCS NCS NCS NCS NCS CS CS CS CS CS EM00@ 75 C ‘ mA ES ude: a a b b b B B B B a a © oO ion oO % Germination bh (o) Cn-D Cn-W 1:50 1:25 1:10 Cn-D Cn-W 1:50 1:25 1:10 NCS NCS NCS NCS NCS CS CS CS CS _ GCS NH98 @ 18/6 C 100 ie B AB A a a a a 80 a a a a % Germination L oO Cn-W 1:50 1:25 1:10 Cn-W_ 1:50 1:25 1:10 NCS NCS NCS NCS CS CS CS CS Fic. 5. Experiment 5 backtransformed germination data, analyzed for each population and incubation temperature separately. Upper case letters indicate significant differ- ences (Tukey’s test, alpha = 0.05) among the three liquid smoke (mQ) concentrations. Lower case letters designate ANOVA result for differences between cold-shocked (CS) and no-shock control (NCS) seeds. Error bars = 1 SD (n = 5). Solid bars are NCS and open bars are CS treatments. Cn-D = dry control; Cn-W = wet control. 120 AcQ OC #8Qm 100 % Germination o oO SYS AAAAANAAWI||]IDH D | 24 82 4 8 ARO) eS AGE 24 8248 4 8 BS (00) Ni, FIG. 6. | | 248248 4 8 SR101 —————_» [Vol. 49 ing the need to understand the effects of fire on plant reproduction. Fire treatments. Smoke applied dry or wet was successful in substantially increasing germination of all dormant seeded populations, sometimes by as much as three to four fold over controls. Dry smoke (S), smoke water (W), and liquid smoke (Q, Qm- 1, Qm-2) all improved germination substantially over controls showing that smoke can break dor- mancy when delivered to seeds in various ways. Because seeds readily imbibed water, the mecha- nism for the effect of smoke on germination in this species may not involve changing the structure of the seed coat cuticle as has been detected for Em- menanthe penduliflora (Egerton-Warburton 1998; Keeley and Fotheringham 1998a). It is possible that ethylene or other components of smoke influence seed germination in California poppy, but this needs further study. Both dry smoke and smoke water contain ethylene (Sutcliffe and Whitehead 1995), which is known to promote germination by several different mechanisms and can sometimes Overcome secondary dormancy or particular tem- perature requirements for germination (Corbineau and Come 1995; Baskin and Baskin 1989). Commercially produced liquid smoke gave the best results and outperformed dry smoke except when applied after chilling. In each case, the seeds responded best to the most natural sequence of events. Any “‘smoke”’ event would likely occur in the dry season before the winter rains or as a result of rain carrying smoke particles from the soil sur- face to the seedbank in late fall or early winter. Results were more erratic for dry smoke than for liquid smoke treatments, possibly because adsorp- tion of dry smoke among experiments is more dif- ficult to control. Other fire treatments were less effective or even inhibited germination. We found no significant ben- efit of charred plant leachate (L), or nitrate (KNO,) on germination. It is possible that charate and NO, I [| | | iW i oe 248248 48 24 H SURO) 1) eS N SOttt{ AAAqNMMHAA_;»4j¥4'i'b'A_'ilidl:! SSSA S8t AadMdAAdMaqCcCM MW MQM MAaAqApQA.A.AaFAajQ_‘_QaiajyaAaga_\_»_o0“ EK... do —~o [HB iy) & & ee) A lee) Experiment 6 backtransformed germination data. Data are arranged by population, treatment, and seed age (in months) along the x-axis. Seeds aged 7.5 weeks are noted as “‘8”’. Data for liquid smoke followed by cold treatment (cQ) were compared with mQ-1 treatment in a separate analysis. Week 4 and week 8 cQ seeds were exposed to 4 weeks at 3.5°C and 2 weeks at 3°C, respectively. Error bars — I SDi@) = 5): 2002] could be more effective if applied before a cold treatment, but this was not tested. Exposure to 85°C for 10 minutes (H and SH) generally reduced ger- mination of viable seeds, decreasing it to about a third of control values. Viability of heat-treated seeds also decreased by an average of 6%, stressing a need for further studies on heat sensitivity of this species. We expect that seeds near the soil surface where temperatures are higher will be largely killed or in- hibited by the heat of fire. In low intensity burns in chaparral, temperatures ranged from about 60°C at a depth of 7-8 cm, about 82°C at 2—3 cm, about 100°C at 1-2 cm deep, to over 180°C at the surface. Also, as fire intensity increased with increase in fuel load, temperatures deep into the seed bank be- came higher (Moreno and Oechel 1991). Higher fire intensities are known to deplete seed banks of other species (Odion 2000). Presumably, prescribed burning of grassland and relatively open California poppy habitats, with less above ground biomass than chaparral, will result in lower fire intensity and heat < 85°C at depths below 2 cm. Our testing at 85°C may adequately represent conditions of a low intensity fire in the upper 2 cm of dry soil where a high density of seeds is expected to occur. Moist soil has higher thermoconductivity which can result in much higher temperatures to greater depths (DeBano et al. 1998). This has implications for the survival of seed banks of temperature sen- sitive species following spring burns over moist soil. In addition, Roche et al. (1998) found much higher germination and survival from smoke treat- ed seedbanks when treated in the dry autumn be- fore winter rains than when treated in winter or spring. Further studies are needed to determine the proportion of seeds killed at different burial depths over a range of fire soil temperatures and moisture levels. Cold treatment and incubation. Many species re- quire cold temperatures to break dormancy or for optimal germination of conditionally dormant seeds (Bewley and Black 1994; Baskin and Baskin 1998), and intra-species differences in response to chilling have been detected in other geographically variable species. For example, in Artemisia tridentata Nutt., which also germinates in winter, among-population differences were found in germination response to different temperatures, with the responses correlat- ing to mean January temperature (Meyer et al. 1990; Meyer and Monsen 1992). In E. californica, populations differed in response to chilling, but it is too early to tell if response correlates with hab- itat. We examined chilling under different moisture levels (dry, moist, wet) because cold winter tem- peratures in California (mid December—February) occur partly while the soil is still dry, and partly after the onset of winter rains. Furthermore, some species have higher seed germination after dry-cold MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 223 storage (Padgett et al. 1999). However, for Califor- nia poppy, compared to no-cold controls, we found a slightly negative effect of wet-cold, no significant benefit of dry-cold, and only a weak effect of moist-cold treatment. Moist-cold resulted in higher germination than no-cold treatment in only six of 13 trials (Table 6). Two reviews of seed dormancy report dormant embryos generally need to be im- bibed before they respond to chilling (Nikolaeva 1969; Baskin and Baskin 1998). It is unlikely that a chilling period longer than 4—8 wk would im- prove germination. In southern California, poppies germinate primarily in January and February, so seeds frequently receive a relatively short period of cold moist exposure, perhaps 2—8 wk, depending on location and onset of rains. In winter 2000, for example, native soils were dry into January in Riv- erside County, yet there was a spectacular bloom that spring. The response of seeds to the various cold treat- ments was, in part, consistent with the dormancy behavior of a winter annual/perennial strategy. Many species in lowland Mediterranean climates germinate and grow in the cool winter rainy season between late fall and late winter. In some of these, prolonged warm summer temperatures or shelf stor- age can break dormancy while many have the abil- ity to germinate at temperatures above about 5°C (Baskin and Baskin 1998). Instead of cold temper- atures necessarily breaking dormancy, very cold temperatures can sometimes send seeds of winter annuals back into dormancy (Baskin and Baskin 1998) or increase degradation of viability (Priestley 1986). In our first two experiments with the lowest temperature cold treatment (2—4°C), dry cold did not cause a decrease in germination, but moist- and wet-cold treatments appeared to either increase dor- mancy of some wild seeds or increase degradation of viable seeds to the extent they lost the ability to germinate. This cold had no effect on domesticated seeds which germinated to nearly 100%. Dry smoke treatment applied before cold had an ame- liorating effect on the lowering of germination (Exp. 2), but smoke treatments applied after cold did not. Cold-shock at 2°C actually decreased ger- mination slightly or had no effect (Exp. 5), a result consistent with a “‘typical’’ winter annual strategy. Treatment at 3—3.5°C did not obviously depress or enhance germination (Exp. 6). Seeds exposed to cold treatments above 4°C began germination in the cold chamber and continued to germinate after moving to warmer incubation chambers (Exp. 3, 4). There was no consistent trend indicating which of the cold temperature treatments improved germi- nation most. Treatment at 4—9°C (mostly 7°C) did not break dormancy in fire controls but did increase germination of smoke-treated seeds. Across exper- iments, germination appeared to improve under cooler incubation temperatures, especially once dormancy was broken with smoke. In addition, the two lower incubation temperatures (7.5 and 18/6°C 224 vs. 22/12°C) resulted in small significant increases in germination (Exp. 5). Lower temperatures are consistent with seed lab testing of California poppy at 15°C (Association of Official Seed Analysts 1981). The temperature range (2—4°C—22/12°C) un- der which perennial, domesticated poppies germi- nated to 100% was quite large, suggesting broad tolerances in the original source populations or se- lection for broad tolerances and loss of conditional dormancy under domestication. Overall, these re- sults show that the effect of cold is subtle as well as population-specific. The results merit further in- vestigation into optimal germination temperatures, how warm storage temperatures (higher than for shelf storage) affect dormancy break, and how ef- fects vary among populations of different life-his- tories (annual vs. perennial) and from different cli- mates. Although California poppy has a small linear embryo, it does not appear to have the morpho- physiological dormancy expected by Martin (1946). If there is such dormancy in California poppy, it is not general to all populations. Even in wild popu- lations with dormant seeds, some seeds appeared to lack physiological dormancy. Extraordinarily high germination of domesticated seeds from commer- cial sources and naturalized garden plants even within two weeks of collection without pretreat- ment, may be due to lost genetic components 1m- portant to germination cycles in natural popula- tions. Most of the fresh seeds collected in 2001 became less dormant as they aged from 2 to 4 and ~8 months. There was some germination by two months suggesting seeds underwent some after-rip- ening by 2 months. Increased germination at 4 mo- old suggests seeds continued to mature in lab stor- age. For all but the Santa Barbara seeds, smoke treatment promoted higher seed germination than controls even at 2 mo, but was even more effective after 4 mo of aging for all populations. Further studies are required to reveal if embryos grow dur- ing dry storage or if their growth follows hydration with or without dormancy breaking smoke treat- ment. The ability of smoke to break dormancy in seeds less than 4 mo of age suggests that smoke from early summer fires may break dormancy pre- maturely, possibly making seeds vulnerable to ger- minating after summer rain if germination could occur at warm temperatures. Because species with morphophysiological dor- mancy sometimes respond to moist-warm temper- ature followed by cold or by GA3 (Nikolaeva 1969; Hidayati et al. 2000), we exposed dormant, aged seeds of FM96 and NH98 to one week of warm- moist stratification at 28°C (A.M.M. and L.F un- published). We chose one week because in southern California’s hot dry summer environment, soil dries out at most within a week of summer storms. Warm stratification did not improve germination over con- - trols even when followed by GA3 (SO0ppm). GA3 treatment improved germination much less than liq- MADRONO [Vol. 49 uid smoke. We plan to test additional combinations of warm stratification followed by cold incubation treatments. Synergistic effects of smoke and cold. We did find a weak synergistic effect of cold treatment in combination with smoke treatments for six popu- lations in an overview of experiments in which smoke treatment was applied before cold treatment (Table 6). Under no-cold, there was consistently higher germination for liquid smoke than no-smoke controls in 15 of 15 trials; under moist-cold, liquid smoke resulted in even higher germination in seven of nine trials (Table 6, Exp. 2—5). Similarly, under no-cold, dry smoke treatment resulted in higher germination than no-smoke controls in 10 of 13 tri- als; under moist-cold, dry-smoke resulted in even higher germination in 10 of 13 trials. This syner- gism suggests smoke is triggering a growth se- quence that must occur before cold temperatures can succeed in promoting germination and may, in part, be substituting for other environmental cues that would normally occur before cold treatment (e.g., warm summer temperatures or exogenous chemicals in the soil). Seed age. Interestingly, young seeds had lower seed dormancy than aged seeds from the same pop- ulations, and dormancy was easier to break than in aged seeds. In young seeds, dry smoke increased germination over the controls, and liquid smoke produced the highest germination of any treatment (62—95% in young seeds vs. O—70% in aged seeds). Viability dropped no more than 6% between young and aged collections from the same sites (viability confirmed with TZ and checked with fluoroscien diacetate methods, unpubl. after Windholm 1972), indicating that older seeds entered a secondary dor- mancy during prolonged shelf storage, or that seeds experienced deterioration in germinability that can- not be detected with chemical viability tests. Ad- ditional cues may be required to break any second- ary dormancy. The ability to enter secondary dor- mancy is important to seasonal cycling of dorman- cy, the building of a seed bank, and presence of a bet-hedging strategy in unpredictable environ- ments. This may be especially important to seeds that germinate in dark so that the seed bank is not exhausted in any particular year. Variation within and among populations. Even though smoke alone or with cold succeeded in in- creasing germination across populations, popula- tions differed substantially in response to those treatments and in baseline dormancy of untreated, shelf-stored seeds. Populations also differed in re- sponse to wet or dry smoke treatments, often in association with annuality or perenniality (Exp. 4). Furthermore, in all wild populations and treatments a fraction of the viable seeds did not germinate. This is important for several reasons: 1) we have not identified a general natural cue that promotes germination of young seeds enmasse in the absence 2002] of fire, or of aged seeds enmasse with or without fire; 2) there are differences among seeds within populations for dormancy and dormancy breaking requirements; and 3) if the variation in response is heritable it can be selected and thus the observed variation in response among populations may be the result of adaptation to local conditions and may be linked to different life-history strategies (Cook 1962; Young and Augspurger 1991). Variation within populations, including changes with seed age, may provide a good bet-hedging strategy. Clearly, the domesticated (Dom and RIV) pop- ulations of California poppy with their non-dormant seeds were very different from all wild populations in our study. Interestingly, some perennial popula- tions from coastal northern California also have no seed dormancy (Cook 1962; Montalvo personal ob- servation, e.g., RMO1). Large differences among wild-collected populations, and between wild and the domesticated populations, show the importance of noting seed source and population traits when doing research or restoration with this species. Our results indicate that results of seed dormancy ex- periments from one or two populations cannot be generalized correctly to the species level. Research on a range of wild populations which have different levels of seed dormancy, including perennial pop- ulations with non-dormant seeds needs to be done before any generalizations can be made about light inhibition of germination and requirements for ger- mination in this species. In pursuit of unknown cues. We have not discov- ered how to break dormancy without smoking seeds. Given that many California poppy seeds ger- minate in nature in the absence of fire, future work should explore the combined effects of seed burial and seasonal changes in temperature on seed ger- mination, with and without smoke. In several Aus- tralian species, prolonged seed burial was found to affect the seed coat by increasing permeability to water and seed coat breakage in ways that facili- tated germination (Tieu and Egerton-Warbuton 2000). In addition, Roche et al. (1997a) found a synergistic effect of lengthy seed burial followed by smoke addition on germination of 60% of over 100 Australian species tested. Smoke treatment by itself approximately doubled germination of seeds, but smoke treatment combined with soil storage quadrupled germination. Seed burial was also found to be important to germination of Dendro- mecon rigida Benth., a fire following species in the Papaveraceae (Keeley and Fotheringham 1998b) that is closely related to the genus Eschscholzia. It is possible that some dormant poppy seeds require a combination of seed burial and changing temper- atures before they can successfully germinate. Al- though exposure to warm summer temperatures breaks dormancy in many winter annuals (Baskin and Baskin 1998), in a recent seed burial study where the soil was dry, as is natural in this region, MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 225 we found that exposure to natural summer and early fall temperatures (July—mid November) did not no- tably improve germination relative to shelf-stored controls (A. Montalvo and C. Koehler unpublished data). Implications for conservation and_ restoration. Given the beneficial effect of smoke treatments on California poppy germination, prescribed fire might prove useful for poppy management as long as heat does not penetrate into the seed bank too deeply. Liquid smoke treatment of the seed bank may sim- ulate the beneficial effects of fire when prescribed burning is not feasible or when heat might unduly affect viability of poppy or other native species. On a cautionary note, evaluation of the differential ef- fects of smoke on other species, both native and exotic, is warranted before using smoke and/or fire aS a management tool. Seeds of exotic and native species may differ in response to smoke treatment and changes in seed coat chemistry may be irre- versible (Roche et al. 1998). If smoke facilitates a non-reversible breaking of seed dormancy for a large fraction of the seed bank, then effects of fre- quent or even occasional burning could last more than one season and could disrupt the selective val- ue of dormancy. The seedbank could be depleted if seeds germinate under conditions detrimental to successful establishment. It is important to weigh the consequences of such negative direct effects of repeated burning on the seed bank, against the ben- eficial effects of killing competitors, or perceived benefit of managing for large flowering displays. Our results can help to develop regionally adapt- ed populations of California poppy for restoration and revegetation projects in southern California, which have in the past met with mixed success. In xeric, non-irrigated locations where native poppies tend to have largely dormant seeds, plantings of domesticated seeds can die out within a few years (A. Montalvo personal observation). Most com- mercially available poppy seeds are non-dormant, perennial, and have resulted from many generations of seed increase, a practice that selects against seed dormancy. The seed industry has avoided dealing in native, dormant seeded populations, in part be- cause it is difficult to break their seed dormancy (Victor Schaff, S&S Seeds personal communica- tion). In the future, the seed industry could smoke- treat dormant native poppy seeds before planting to avoid selecting against seed dormancy. Given the evidence for locally adaptive differences and home site advantage in many other plant species (for re- view see Langlet 1971; Montalvo and Ellstrand 2000), and the sometimes adverse effects of hy- bridization among genetically differentiated popu- lations (Millar and Libby 1989; Knapp and Rice 1994: Montalvo et al. 1997, Montalvo and Ellstrand 2001), the use of local poppy seeds for restoration and reseeding would likely increase the long-term success of planted populations. 226 ACKNOWLEDGMENTS We thank John Crossman, Gordon Fox, Robert Noll of Noll seeds and Victor Schaff of S&S Seeds for generous contributions of native seed collections, two anonymous reviewers for suggestions to improve the manuscript, Hi- lary Wall for unpublished information about effects of liq- uid smoke on other local native plants, L. Egerton-War- burton for discussions and information about seed ger- mination biology and seed viability testing, undergraduate students M. Sripracha, J. Terrell, and M. Blatt who helped with seed processing and scoring, the Antelope Valley California Poppy Reserve and Western Riverside Multi- Species Reserve for permission to collect seeds from re- serve populations, Regen of Glasgow Kentucky and Gray- son Australia of Bayswater, Victoria for information and samples of Regen 2000 liquid smoke, and California De- partment of Parks and Recreation (Agreement No. 67834) and Metropolitan Water District of Southern California (Agreement No. 4602) for financial support. LITERATURE CITED ASSOCIATION OF OFFICIAL SEED ANALYSTS. 1981. Rules for testing seeds. Journal of Seed Technology 6:i—iv, 1— IZ. BASKIN, C. C. AND J. M. BASKIN. 1998. Seeds: ecology, biogeography and evolution of dormancy and ger- mination. Academic Press, San Diego, CA. BEAN, L. J. AND H. W. LAwTon. 1973. 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MADRONO, Vol. 49, No. 4, pp. 228-236, 2002 TEMPERATURE LIMITATIONS FOR CULTIVATION OF EDIBLE CACTI IN CALIFORNIA PARK S. NOBEL!, ERICK DE LA BARRERA, DAVID W. BEILMAN, JENNIFER H. DOHERTY, AND BRIAN R. ZUTTA Department of Organismic Biology, Ecology, and Evolution, University of California, Los Angeles, CA 90095-1606 ABSTRACT Hylocereus undatus (a hemiepiphyte) and Opuntia ficus-indica (“prickly pear’’) are cultivated world- wide as specialty fruit and vegetable crops, so the role of temperature in determining regions suitable for commercial growth of these cacti was investigated for California, the leading agricultural state in the United States. Air temperatures below —2.5°C and above 45°C are lethal for H. undatus compared with below —10°C and above 65°C for O. ficus-indica, demonstrating the latter’s greater tolerance of extreme temperatures. Mean nighttime air temperatures influence net CO, uptake for these Crassulacean acid metabolism species, optimal uptake occurring at 20°C for H. undatus and at 14°C for O. ficus-indica. Extreme air temperatures over a 30-year period for 326 weather stations and mean nighttime temperatures for 259 stations were mapped to identify where these species could be cultivated. Only 2% of the state’s total area avoided temperatures lethal to H. undatus, mostly along the southern coast. In contrast, 36% of the state’s area was possible for O. ficus-indica, exclusion occurring in mountainous regions. A Tem- perature Index (net CO, uptake over 24-hour periods at a particular temperature divided by uptake at the optimal temperature) was also utilized to evaluate a region’s suitability for growing these cacti. The Temperature Index was below 0.7 for 59% of the weather stations for H. undatus but for only 16% for O. ficus-indica. In the regions where lethal extreme temperatures did not occur, the Temperature Index averaged more than 0.8 for both species. Use of a Temperature Index based on net CO, uptake together with extreme temperature events can help evaluate regions for cultivating cacti with edible fruits or other new crops. Key Words: CO, uptake, Crassulacean acid metabolism, fruit, Hylocereus undatus, Opuntia ficus-indica. As a result of its geology, topography, and cli- mate, California is the most biodiverse state in the United States, having nearly 6000 native species of vascular plants (Hickman 1993). Such variety also permits the production of about 350 agricultural commodities that generate $30 billion in revenue per year, more than for any other state, such com- modities being responsible for 10% of the jobs in California (California Department of Food and Ag- riculture 2001). Much of the agricultural economy derives from intensively managed specialty crops— e.g., minor crops such as grapes, tomatoes, straw- berries, lettuce, and flowers generate about 20% of the agricultural sector’s annual revenue (California Department of Food and Agriculture 2001). In this regard, considerable interest exists among both government officials and also producers to develop new specialty crops, such as the cacti with edible fruits considered here. Given their potentially high productivity and tol- erance of high temperatures (Nobel 1988), cacti have become important crops in arid and semiarid regions worldwide (Barbera 1995). The prickly pear cactus Opuntia ficus-indica (L.) Miller is cul- tivated on over one million hectares in more than 30 countries primarily for fodder but also as a fruit ' Author for correspondence, e-mail: psnobel@biology. ucla.edu crop (“‘cactus pears’’) and on a limited scale as a vegetable crop (“‘nopalitos’’; Nobel 2000). Second in importance among edible cacti are hemiepiphy- tes in the genera Hylocereus and Selenicereus, whose fruits are commonly referred to as “‘pitahay- as,’ which are cultivated in 20 countries, with par- ticularly noteworthy success in Southeast Asia (Nerd et al. 2002; Nobel and De la Barrera 2002). Cacti have been cultivated in California since the eighteenth century, when O. ficus-indica was plant- ed around the Spanish missions along the coast for its fruit and its mucilage, which was utilized as a binding material for adobe bricks (McLeod 1975). In the early 1900s, Luther Burbank, who pioneered several specialty crops, developed a “‘spineless”’ variety of O. ficus-indica (Savio 1989). The D’ Arrigo Brothers have plantations of spineless O. ficus-indica in the Santa Clara Valley (also known as the Salinas Valley) near Gilroy that were estab- lished in the 1920s for fruit (Curtis 1977), and var- ious companies in southern California are currently exploring the pitahaya market (Savio 1989; Valdi- via 2000; P.S. Nobel personal observation). Most gas exchange for cacti occurs at night when air temperatures are lower than during the daytime, a characteristic of the Crassulacean acid metabo- lism (CAM) photosynthetic pathway. This physio- logical strategy, which has evolved for species in over 30 vascular plant families, reduces transpira- 2002] tional water loss (Nobel 1999; Taiz and Zeiger 2002). Optimal nocturnal air temperatures for net CO, uptake by CAM plants are generally from 10 to 20°C (Nobel 1988). Extreme temperatures limit CO, uptake and also can damage cacti, with epi- sodic freezing temperatures generally being more limiting to plant distribution than high tempera- tures. For instance, based on the uptake of a vital stain into the central vacuoles, the photosynthetic cells (chlorenchyma) of O. ficus-indica have 50% mortality (LT;,) at the extremely high temperature of 62.4°C for plants kept at day/night air tempera- tures of 40/30°C and at 66.6°C for plants kept at 50/40°C, indicating that O. ficus-indica has a high- temperature acclimation (hardening) of 4.2°C per 10°C increase in air temperature (Nobel 1988). The LT;, for low-temperature tolerance of O. ficus- indica is —7.7°C for plants kept at day/night air temperatures of 20/10°C, decreasing to —8.8°C for plants kept at 10/0°C, indicating a low-temperature acclimation of 1.1°C for a temperature decrease of 10°C (Nobel 1988). Hylocereus undatus (Haworth) Britton & Rose, which is native to neotropical for- ests (Britton and Rose 1963; Backeberg 1966; Barthlott and Hunt 1993) where temperatures tend to be warm and vary little over the course of a year (Croat 1978; Liittge 1997), shows a high-tempera- ture acclimation of only 1.4°C per 10°C increment in air temperature, as LT;, occurs at 54.0°C for plants kept at day/night air temperatures of 25/15°C and at 55.4°C for plants at 35/25°C; furthermore, plants kept at 40/30°C develop stem tissue necrosis, leading to death after 19 weeks (Nobel and De la Barrera 2002). Because the responses of H. undatus to freezing temperatures have not been reported, one of the objectives of the present research was to determine its low-temperature tolerance and its low-temperature acclimation. An Environmental Productivity Index (EPI), which can help to evaluate the feasibility for ex- panding the area of cultivation of crops such as cacti, indicates the primary influence of water, tem- perature, and light on net CO, uptake and hence biomass productivity of plants (Nobel 1988, 1999). EPI is defined as the Water Index X the Tempera- ture Index X the PPF Index (PPF refers to the pho- tosynthetic photon flux, composed of wavelengths of light from 400 to 700 nm), where each compo- nent index ranges from 0.00, when that environ- mental factor eliminates net CO, uptake, to 1.00, when that factor is optimal for net CO, uptake (No- bel 1988). EPI ignores secondary interactions, such as the different response to PPF when temperature is limiting, but in any case net CO, uptake is gen- erally low under such conditions. The individual indices are determined in the laboratory under con- trolled conditions over 24-hour periods by varying the environmental parameter to be studied, while keeping the other factors constant at optimal values. EPI can then be calculated under field environmen- tal conditions, as has been done to predict quanti- NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI 229 — Hylocereus undatus —— Opuntia ficus-indica Daily net CO, uptake per unit stem area (fraction of maximum) eee L Spee l 5 10 15 20 25 30 35 Mean nighttime air temperature (°C) Fic. 1. Responses of total daily net CO, uptake by Hy- locereus undatus and Opuntia ficus-indica to mean night- time air temperatures. Plants were maintained for 7 to 14 days at a particular temperature before measurement. Data for H. undatus are from Raveh et al. (1995) and Nobel and De la Barrera (2002) and for O. ficus-indica are from Nobel (1988) and Nobel and Bobich (2002). tatively the growth of Agave deserti along an ele- vational gradient (Nobel 1984) and to establish new plantations of O. ficus-indica in Chile (P. S. Nobel personal observation). For agricultural purposes, the Water Index can be increased to 1.0 by irriga- tion, and the PPF Index can be manipulated by varying the spacing between plants. However, it is difficult to control temperatures in the field. Deter- mining the Temperature Index may therefore help producers decide the suitability of a particular re- gion for growing a certain crop. In this regard, net CO, uptake is optimal at a mean nighttime temper- ature of 20°C for H. undatus and 14°C for O. ficus- indica (Fig. 1). Also, H. undatus has a narrower temperature range than does O. ficus-indica within which the Temperature Index is above 0.5 (9 to 26°C versus 2 to 26°C) and above 0.8 (14 to 23°C versus 6 to 20°C). The present research uses the previously measured Temperature Indices for H. undatus and O. ficus-indica (Fig. 1), the low and the high temperature tolerances for both species, and climatic data to evaluate potential regions for their cultivation in California. This approach used for cacti here can also serve as a model for assess- ing the cultivation potential of other plant species. MATERIALS AND METHODS Temperature tolerances. The low-temperature tolerance has already been determined for Opuntia ficus-indica, as have the high-temperature toler- ance for it and Hylocereus undatus (Nobel 1982, 1988; Nobel and De la Barrera 2002), using the vacuolar uptake of neutral red (3-amino-7-dimeth- ylamino-2-methylphenazine hydrochloride) fol- lowing exposure for 60 min to a particular extreme temperature (Onwueme 1979; Didden-Zopfy and Nobel 1982; Nobel et al. 1995). Hence, the low- temperature tolerance of H. undatus was similarly 230 determined here using the neutral red assay. For this purpose, five plants approximately 45 cm in shoot length were obtained from the Cactus Trad- ing Company (Jamul, CA). They were grown in each of two Conviron E-15 environmental cham- bers (Controlled Environments, Pembina, ND) with day/night air temperatures of 20/10°C or 30/ 20°C for 4 weeks with weekly application of 0.2- strength Hoagland’s solution supplemented with micronutrients and a total daily PPF of 16 mol m * day-!, environmental conditions that are near the optimal for H. undatus (Raveh et al. 1995; Nobel and De la Barrera 2002). Low temperatures at 1 to 2°C intervals decreas- ing from 4°C were obtained in an ULT-80 ultra- low-temperature freezer (Rheem Manufacturing, West Columbia, SC). Approximately 1.5 g of stems were removed with a scalpel, placed in contact with a copper-constantan thermocouple 0.51 mm in di- ameter, and wrapped in aluminum foil to prevent desiccation; the samples were then cooled at 5°C hr~', similar to stem cooling rates observed in the field (Nobel 1988; Nobel et al. 1995). After expo- sure to a particular temperature for 60 min, the samples were sliced into sections approximately 700 wm thick using razor blades and then placed in 0.2% (w/w) neutral red for 90 min for stain up- take, which occurs for the vacuoles of living cells only and indicates membrane integrity (Onwueme 1979; Nobel et al. 1995). The tissue samples were then placed for 10 min in 0.25 M potassium phos- phate buffer (pH 7.8) at 25°C followed by 24 hours at 6°C in distilled water to help remove excess stain and hence to sharpen the images, after which ap- proximately 130 intact cells per sample were ex- amined at 100X using a BH-2 phase-contrast mi- croscope (Olympus, Lake Success, NY) to check for stained (living) versus unstained cells. The low temperature treatment that halved stain uptake from the maximum occurring at 4°C (LT.,), a reliable test for predicting eventual tissue necrosis (Didden- Zopfy and Nobel 1982; Smith et al. 1984; Nobel et al. 1995), was determined graphically under each condition. Low-temperature acclimation was ana- lyzed by comparing LT;, for plants at day/night air temperatures of 20/10°C versus 30/20°C using an unpaired Student t-test. Extreme temperature limitation. For cacti in na- ture as well as under cultivation, infrequent freez- ing episodes can be severely limiting (Steenbergh and Lowe 1976; Russell and Felker 1987; Nobel 1988). Moreover, fruit production by H. undatus and O. ficus-indica can occur two years after the establishment of a plantation, but approximately 10 years may be necessary to obtain optimal yields and an even longer period for appropriate return on the initial investment (Mizrahi and Nerd 1999; In- glese et al. 2000; Nerd et al. 2002). Also, the fruit- ing potential of O. ficus-indica tends to decrease after 25 to 30 years (Inglese et al. 2002). Thus, MADRONO [Vol. 49 instead of using annual minimum temperatures, the lowest air temperatures recorded between 1961 and 1990 (the most recent period summarized with cli- mate normals) at 326 California weather stations were obtained from the Climate Atlas of the Con- tiguous United States (National Climatic Data Cen- ter 1995). Similarly, the highest air temperatures were obtained for the 318 weather stations with suitable records for the same period. The tempera- tures were converted from Farenheit to Celsius and then ranked in 2.5°C intervals for low-temperature extremes or 5°C intervals for high-temperature ex- tremes. In addition, weather station data were in- terpolated, correcting for elevation using a lapse rate of 6°C per km (Nobel 1999), to identify the areas with record minimum temperatures below —10°C or above —2.5°C as well as record maxi- mum temperatures above 45°C over the 30-year pe- riod (1961-1990). The resulting low-temperature and high-temperature maps were created in Arc- View 3.1 (ESRI, Redlands, CA). Temperature Index. Daily minimum tempera- tures averaged over each month for the California weather stations from 1961 to 1990 and then ay- eraged over the 30 years were also obtained from the Climate Atlas of the Contiguous United States (National Climatic Data Center 1995). Nighttime mean air temperatures were estimated by adding 3°C to the average minimum temperature (Nobel 1988) recorded at each of the 259 weather stations with sufficient records. The Temperature Index for total daily net CO, uptake per unit stem area for H. undatus and O. ficus-indica was then determined for each month using the known temperature re- sponses for these two species (Fig. 1). The twelve monthly values were averaged to obtain an annual Temperature Index for each weather station and maps were created in ArcView 3.1. RESULTS Low-temperature tolerance for Hylocereus un- datus. Neutral red accumulation in chlorenchyma cells of Hylocereus undatus decreased as the treat- ment temperature was lowered below O0°C (Fig. 2). For H. undatus growing at day/night air tempera- tures of 30/20°C, the percentage of cells taking up the vital stain was halved (LT;,) at —1.31 + 0.04°C. The LT... for H. undatus acclimated to day/night air temperatures of 20/10°C was —1.55 + 0.07°C. This species thus displayed a small, yet significant, low- temperature acclimation (hardening) of 0.24 = 0.08°C per 10°C decrease in temperature (t = 2.98, P < 0.01, df = 8). Extreme temperature limitations. During the 30- year period considered (1961-1990), 40% of the 326 weather stations had temperatures below —10°C, corresponding to 64% of the area of Cali- fornia, mainly in regions at high elevations in the Sierra Nevada, Coast and Diablo ranges, and the San Bernardino Mountains, as well as in northern 2002] oy oO oO T ies) oOo T =O SANE 7 —A— 20/10°C (% of maximum) Chlorenchyma cells taking up stain 3 2 -1 0 1 2 3 4 Temperature (°C) Fic. 2. Influence of day/night air temperatures of 20/ 10°C and 30/20°C on the low-temperature tolerance of H. undatus. Uptake of neutral red was determined for pieces of chlorenchyma incubated at a particular treatment tem- perature for 60 min. Data are means = SE (n = 5 plants). >0 0 to-2.5 -2.5 to -5 -5 to-7.5 -7.5 to -10 < -10 *ODQA0 70 ® Fic. 3. NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI 254 California (Fig. 3A). Regions with extreme mini- mum temperatures from —5 to —10°C (49% of the Stations) are concentrated along the coast, through the Central Valley, and in southern California. Re- gions that were never below —5°C (11% of the sta- tions) are largely restricted to thin coastal regions in the San Francisco Bay area, the Channel Islands, and in southern California from Ventura to San Di- ego counties, in addition to lower inland elevations in southern California. Regions with extreme min- imum temperatures above —2.5°C (7% of the sta- tions), representing only 2% of the state’s area, are located along the coast in Ventura, Los Angeles, Orange, and San Diego counties plus one station in the San Francisco Bay area (Fig. 3B). Only one station (at the University of California, Los Ange- les) remained above 0°C during the period consid- ered (1961-1990). During the same 30-year period, five weather sta- tions had maximum temperatures above 50°C 20 to 25 25 to 30 30 to 35 35 to 40 40 to 45 45 to 50 > 50 3 ee et ar ef (| Temperature extremes at California weather stations from 1961—1990: (A) record minimum temperatures and (B) record maximum temperatures. Letters indicate specific ranges of extreme temperatures, e.g., c corresponds to a record minimum temperature between —2.5°C and —5°C. 232. 0.0 to 0.1 0.1 to 0.2 0.2 to 0.3 0.3 to 0.4 0.4 to 0.5 0.5 to 0.6 0.6 to 0.7 0.7 to 0.8 0.8 to 0.9 0.9 to 1.0 0 1 2 3 4 5 6 la 8 9 Fic. 4. MADRONO [Vol. 49 Annual Temperature Index values at California weather stations averaged over 30 years (1961—1990) for (A) H. undatus and (B) O. ficus-indica. Each number represents the influence of temperature on the fraction of maximal total daily net CO, uptake averaged over the year, as calculated from monthly mean nighttime air temperatures, for each weather station. Numbers indicate specific ranges of the annual Temperature Index, e.g., 7 corresponds to 0.7 to 0.8. (1.6% of the 318 stations), with the hottest site (53°C) in Death Valley (Fig. 3B). Maximum tem- peratures from 45 to 50°C occurred at 26% of the weather stations, concentrated at lower elevations in the Mojave Desert and Death Valley, along the Central Valley, and in inland southern California. Approximately 40% of the state’s area had temper- atures above 45°C during the period considered. Regions with maximum temperatures from 40 to 45°C for the 30-year period considered (53% of the stations) are situated at higher elevations in north- ern California, in the Central Valley, and along the coast from the San Francisco Bay area through San Diego County. Only 20% of the stations, distributed in coastal regions or at high mountain elevations, recorded maximum temperatures below 40°C (Fig. 3B). Temperature Index. For H. undatus, the annual Temperature Index (TI) was below 0.5 for 23% of the 259 weather stations, mostly those at high ele- vations, especially in northern and inland California (Fig. 4A). An annual TI of 0.5 to 0.7 occurred for 36% of the stations, most occurring in the Central and Imperial valleys. An annual TI from 0.7 to 0.8 was restricted to Sacramento, the San Francisco Bay area, and the southern California coast (33% of the stations; Fig. 4A). A TI of 0.8 to 0.9 was found only in Los Angeles, Orange, and San Diego counties as well as in Death Valley (8% of the sta- tions). Compared with H. undatus, the annual TI tended to be higher for Opuntia ficus-indica, only 4% of the weather stations having annual values below 0.5 (Fig. 4B). An annual TI of 0.5 to 0.7 occurred for 2002] 12% of the stations, mostly in inland regions, es- pecially for northeastern California. Moderately high TI, from 0.7 to 0.8 (18% of the stations) and from 0.8 to 0.9 (32% of the stations), were located throughout California, except at high elevations in the Sierra Nevada. Annual TI values above 0.9 for O. ficus-indica occurred along the coast from the Oregon border south to Ventura County as well as in various inland regions, such as in the San Fran- cisco Bay area and the Los Angeles basin (34% of the stations; Fig. 4B). DISCUSSION Twenty-three species of cacti have been exam- ined for tolerance to extreme temperatures (Nobel 1982, 1988; Smith et al. 1984), which are important in determining natural distributions and potential regions for cultivation. The least freezing tolerant of these species, Opuntia ramosissima, is native to the deserts of the southwestern United States and northern Mexico and has an LT;, (temperature that kills half of the cells compared with the control) of —4.4°C when maintained at day/night air tempera- tures of 10/0°C (Nobel 1982). Hylocereus undatus was even less freezing tolerant, with an LT,, of —1.6°C when maintained at 20/10°C. Taking into consideration its relatively small low-temperature acclimation of 0.2°C per 10°C decrease in air tem- perature observed here, the LT;, for H. undatus is only —1.8°C at 10/0°C, indicating that it is extreme- ly sensitive to freezing temperatures. In addition, H. undatus is not as tolerant of high temperatures as are the other cactus species examined (Nobel 1988) and also exhibits little high-temperature ac- climation (Nobel and De la Barrera 2002). Indeed, acclimation is the key to tolerating extreme tem- peratures, and only one cactus species (Ferocactus covillei) has less low-temperature acclimation and none has less high-temperature acclimation than does H. undatus (Nobel 1988). In this regard, H. undatus is native to neotropical forests with mod- erate and rather stable temperatures (Britton and Rose 1963; Backeberg 1966; Croat 1978; Barthlott and Hunt 1993; Liittge 1997) and apparently is not genetically or physiologically capable of apprecia- ble acclimation to low or to high temperatures, al- though further studies are necessary to understand its intraspecific variation. LT.) is used for its ease of measurement and be- cause it is often the temperature where stem dam- age becomes visible, although cacti generally do not die until the cellular uptake of a vital stain is reduced to zero. As assessed by neutral red stain- ing, stem death of most cacti occurs approximately 4°C below the low-temperature LT,, and 4°C above the high-temperature LT., (Nobel et al. 1986; Nobel 1988). Moreover, LT., refers to tissue temperatures when damage occurs, not air temperatures, which can differ significantly. On clear nights, tempera- tures of cactus stems can be a few degrees Celsius NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI i) Oo oS) below air temperature due to transpirational cooling and especially net heat loss by infrared (longwave) radiation (Nobel 1988, 1999). Indeed, radiation frosts, when the tissue achieves freezing tempera- tures with air temperatures above O°C, are a severe agricultural problem in California, especially for the citrus industry (Pehrson 1984), and affect the suitability of a site for cactus cultivation. Freezing temperatures can cause extracellular ice crystal for- mation in cacti, which draws water out of the cells and can lead to irreversible damage (Burke et al. 1976; Nobel 1982, 1988). For regions that experi- ence infrequent damaging or even lethal low tem- peratures, freeze-protection methods, such as shade cloth, heaters, and overhead irrigation, can mitigate freezing damage to perennial plants such as H. un- datus and O. ficus-indica (Pehrson 1984; Perry 1998). Differences between air and tissue temperatures can be even larger during the day than at night, depending on stem orientation relative to solar ir- radiation and stem massiveness (Nobel 1988). Tis- sue temperatures of the relatively thin stems of H. undatus are not expected to rise more than | to 2°C above air temperature, especially in its typically shaded habitat (Nobel and De la Barrera 2002). In contrast, stem temperatures for O. ficus-indica can be more than 15°C above air temperatures (Wallace and Clum 1938; Konis 1950). Extremely high tem- peratures can denature proteins, degrade cell mem- branes, and disrupt metabolism in general (Nobel 1988; Srinivasan et al. 1996; Taiz and Zeiger 2002). For H. undatus, daytime temperatures of 45°C can reduce flower and hence fruit production (Mizrahi and Nerd 1999). Shade cloth has been used to ame- liorate the effects of high temperatures (and high PPP) on A. undatus growing in Israel (Raveh et al. 1998). In contrast, stems of O. ficus-indica usually are not damaged until air temperatures exceed 65°C and it can even tolerate 60 min at 70°C (Nobel et al. 1986; Nobel 1988), so high temperatures should not be a limiting factor for cultivation of this cactus in California. Hylocereus undatus can be grown in regions with extreme temperatures above —2.5°C and be- low 45°C (Fig. 5A), which occur for only 2% of the state’s area. On the other hand, O. ficus-indica is excluded only from regions of California where the minimum temperature is below —10°C (Fig. 5B) and can be grown in 36% of the state’s area. The climate of California, which renders most of the state too cold for maximal net CO, uptake by H. undatus, resulted in a lower annual Temperature Index (TI) for it, averaging 0.57 throughout the state compared to 0.82 for O. ficus-indica. The low- er average annual TI for H. undatus reflects both its relatively high optimal mean nighttime temper- ature for net CO, uptake and also the more rapid decrease in net CO, uptake above and below the optimal value than is the case for O. ficus-indica. In the regions where the cacti can be cultivated be- 234 MADRONO [Vol. 49 Fic. 5. Annual Temperature Index values (from Fig. 4) for the weather stations with extreme temperatures within the tolerable ranges for (A) H. undatus (extreme temperatures between —2.5°C and 45°C) and (B) O. ficus-indica (minimum temperature of —10°C and no maximum temperature). cause of lack of lethal extreme temperatures (Fig. 5), the annual TI averages 0.83 for H. undatus and 0.90 for O. ficus-indica, both high values, indicat- ing that the nighttime temperatures for regions within the extreme temperature limits are condu- cive to substantial net CO, uptake by these two spe- cies. The similarly high annual TI estimated for O. ficus-indica in areas suitable for cultivation (and in the entire state) reflect the wide range of nighttime temperatures at which this species can be grown successfully. Nevertheless, the exclusion of 64% of California’s area indicates that even single extreme low-temperature events can greatly damage plan- tations of O. ficus-indica (Russell and Felker 1987; Nobel 1988). Frost damage can be avoided with appropriate agricultural practices (Pehrson 1984; Perry 1998) or by the utilization of cold-tolerant cultivars of Opuntia, a genus with considerable ge- netic diversity as well as a long history of agricul- tural selection (Russell and Felker 1987; Parish and Felker 1997; Casas and Barbera 2002). The Temperature Index is the least manageable of the three components of the Environmental Pro- ductivity Index (EPI) in an agricultural setting, which is the reason for its consideration in the pre- sent study. The relationship between EPI, a predic- tor of net CO, uptake, and fruit production has not been investigated for H. undatus or O. ficus-indica, but EPI closely predicts leaf production for Agave tequilana (Nobel and Valenzuela 1987) and cladode production for O. ficus-indica (Nobel 1988) under cultivation. Besides the present focus on fruit crops and the use of young cladodes of O. ficus-indica as a vegetable, most cultivation of cacti worldwide is dedicated to fodder production (Nobel 2000), due to ease of management without irrigation or fertil- izer application and an acceptable protein content of 5 to 8% on a dry mass basis (Nobel 1988; Pi- 2002] mienta Barrios 1990). Such fodder could be used in California as an input to the state’s livestock and poultry sector, which is responsible for 10% of the state’s agricultural revenue (California Department of Food and Agriculture 2001). In any case, the market for cactus fruits, which had been restricted for cactus pears in Mexico and southern Italy as well as for pitahayas in southeastern Asia, has re- cently expanded globally. The establishment of new plantations of these and other edible cacti in Cali- fornia, for domestic consumption by ethnic groups who traditionally consume cacti plus others who are developing a taste for these exotic fruits plus ex- portation, could contribute to the diversification of revenue production for this leading agricultural State. ACKNOWLEDGMENTS We thank Kevin Coniff for providing the plants of Hy- locereus undatus and the UCLA-Ben Gurion University Program of Cooperation for financial support through the generous gift of Dr. Sol Leshin and the continuing dedi- cation to such projects by Professor Samuel Aroni. LITERATURE CITED BACKEBERG, C. 1966. Das Kakteenlexikon. Gustav Fisher, Jena, Germany. : BARBERA, G. 1995. History, economic and agro-ecological importance. Pp. I-11 in G. Barbera, P. Inglese, and E. Pimienta-Barrios (eds.), Agro-ecology, cultivation and uses of cactus pear. FAO Plant Production and Protection Paper 132. FAO, Rome, Italy. BARTHLOTT, W. AND D. R. Hunt. 1993. Cactaceae. Pp. 161—196 in K. Kubitzki (ed.), The families and gen- era of vascular plants, Vol. 2. Springer-Verlag, Berlin, Germany. BriTTon, N. L. AND J. N. Rose. 1963. The Cactaceae: descriptions and illustrations of plants of the cactus family, Vol. Il. Dover, New York, NY. BurKgE, M. J., L. V. Gusta, H. A. QUAMME, C. J. WEISER, AND P. H. Li. 1976. Freezing and injury in plants. Annual Review of Plant Physiology 27:507—528. CALIFORNIA DEPARTMENT OF FOOD AND AGRICULTURE. 2001. California Department of Food & Agriculture Resource Directory 2000. California Department of Food and Agriculture, Sacramento, CA. Casas, A. AND G. BARBERA. 2002. Mesoamerican domes- tication and diffussion. Pp. 143-162 in P. S. Nobel (ed.), Cacti: biology and uses. University of Califor- nia Press, Berkeley, CA. Croat, T. B. 1978. Flora of Barro Colorado Island. Stan- ford University Press, Stanford, CA. Curtis, J. R. 1977. Prickly pear farming in the Santa Clara Valley, California. Economic Botany 31:175—179. DIDDEN-Zopry B. AND P. S. NoBEL. 1982. High tempera- ture tolerance and heat acclimation of Opuntia bige- lovii. Oecologia 52:176—180. HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. INGLESE, P., EF BASILE, AND M. SCHIRRA. 2002. Cactus pear fruit production. Pp. 163-183 in P. S. Nobel (ed.), Cacti: biology and uses. University. of California Press, Berkeley, CA. NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI 235 Konis, E. 1950. On the temperature of Opuntia joints. Palestine Journal of Botany, Jerusalem Series 5:46— SDs LurtcE, U. 1997. Physiological ecology of tropical plants. Springer-Verlag, Berlin, Germany. McLeop, M. G. 1975. A new hybrid fleshy-fruited prick- ly-pear in California. Madrono 23:96—98. MIZRAHI, Y. AND A. NERD. 1999. Climbing and columnar cacti: new arid land fruit crops. Pp. 358-366 in J. Janick (ed.), Perspectives on new crops and new uses. ASHS Press, Alexandria, VA. NATIONAL CLIMATIC DATA CENTER. 1995. Climate atlas of the contiguous United States. National Climatic Data Center, Asheville, NC. NERD, A., N. TEL-ZUR, AND Y. MIZRAHI. 2002. Fruits of vine and columnar cacti. Pp. 185-197 in P. S. Nobel (ed.), Cacti: biology and uses. University of Califor- nia Press, Berkeley, CA. NoBEL, P. S. 1982. Low-temperature tolerance and cold hardening of cacti. Ecology 63:1650—1656. . 1984. Productivity of Agave deserti: Measure- ments by dry weight and monthly prediction using physiological responses to environmental parameters. Oecologia 64:1-—7. . 1988. Environmental biology of agaves and cacti. Cambridge University Press, New York, NY. . 1999. Physicochemical and environmental plant physiology, 2nd ed. Academic Press, San Diego, CA. . 2000. Crop ecosystem responses to climatic change: Crassulacean acid metabolism crops. Pp. 315-331 in K. R. Reddy and H. E Hodges (eds.), Climate change and global crop productivity. CABI Publishing, New York, NY. AND E. G. BosicH. 2002. Environmental biology. Pp. 57-74 in P. S. Nobel (ed.), Cacti: biology and uses. University of California Press, Berkeley, CA. AND E. DE LA BARRERA. 2002. High temperatures and net CO, uptake, growth, and stem damage for the hemiepiphytic cactus Hylocereus undatus. Biotropica 34:225-—231. AND A. G. VALENZUELA. 1987. Environmental re- sponses and productivity of the CAM plant, Agave tequilana W. Agricultural and Forest Meteorology 39: 319-334. , G. N. GELLER, S. C. KEE, AND A. D. ZIMMERMAN. 1986. Temperatures and thermal tolerances for cacti exposed to high temperatures near the soil surface. Plant, Cell and Environment 9:279—287. , N. WANG, R. A. BALSAMO, M. E. LOoIk, AND M. A. HAWKE. 1995. Low-temperature tolerance and ac- climation of Opuntia spp. after injecting glucose or methylglucose. International Journal of Plant Scien- ces 156:496—504. ONWUEME, I. C. 1979. Rapid, plant-conserving estimation of heat tolerance in plants. Journal of Agricultural Science, Cambridge 92:527—536. PARISH, J. AND P. FELKER. 1997. Fruit quality and produc- tion of cactus pear (Opuntia spp.) fruit clones selected for increased frost hardiness. Journal of Arid Envi- ronments 37:123—143. PEHRSON, J. 1984. Pointers on planning a citrus frost pro- tection program. Citrograph 69:263-—265. PERRY, K. B. 1998. Basics of frost and freeze protection for horticultural crops. HortTechnology 8:10—15. PIMIENTA BARRIOS, E. 1990. El nopal tunero. Universidad de Guadalajara, Guadalajara, Mexico. RAVEH, E., M. GERSANI, AND P. S. NOBEL. 1995. CO, up- take and fluorescence responses for a shade-tolerant 236 cactus Hylocereus undatus under current and doubled CO, concentrations. Physiologia Plantarum 93:505— Sule , A. NERD, AND Y. MIZRAHI. 1998. Responses of two hemiepiphytic fruit-crop cacti to different degrees of shade. Scientia Horticulturae 73:151—164. RUSSELL, C. AND P. FELKER. 1987. Comparative cold-har- diness of Opuntia spp. and cultivars grown for fruit, vegetable and fodder production. Journal of Horti- cultural Science 62:545—550. SAvio, Y. 1989. Prickly pear cactus. Family Farm Series, Small Farm Center, University of California. Davis, CA. SMITH, S. D., B. DIDDEN-ZopFy, AND P. S. NOBEL. 1984. High temperature responses of North American cacti. Ecology 65:643—651. MADRONO [Vol. 49 SRINIVASAN, A., H. TAKEDA, AND T. SENBOKU. 1996. Heat tolerance in food legumes as evaluated by cell mem- brane thermostability and chlorophyll fluorescence techniques. Euphytica 88:35—45. STEENBERGH, W.E AND C.H. Lowe. 1976. Ecology of the saguaro: I. The role of freezing weather in a warm- desert population. Pp. 49—92 in Research in the parks, National Park Service Symposium Series 1, U.S. Government Printing Office, Washington, DC. Taiz, L. AND E. ZEIGER. 2002. Plant physiology, 3rd ed. Sinauer Associates, Sunderland, MA. VALDIVIA, E. 2000. Pitahaya: A fruit for the diligent. Fruit Gardener 32:12-13. WALLACE, R. H. AND H. H. CLum. 1938. Leaf tempera- tures. American Journal of Botany 25:83—97. MADRONO, Vol. 49, No. 4, pp. 237-255, 2002 SIX NEW SPECIES AND TAXONOMIC REVISIONS IN MEXICAN GAUDICHAUDIA (MALPIGHIACEAE) STEVEN L. JESSUP Department of Biology, Southern Oregon University, Ashland, OR 97520 Jessup @ sou.edu ABSTRACT Six new species in the genus Gaudichaudia are described and sectional taxonomy in the genus is revised in accordance with findings from molecular analysis of chloroplast and nuclear genomes. Gau- dichaudia cycloptera, G. chasei, and G. mcvaughii are removed from section Gaudichaudia and placed in section Cyclopterys. Gaudichaudia krusei and G. subverticillata are removed from section Gaudi- chaudia and placed in section Archaeopterys. Gaudichaudia hirtella, comb. nov. is placed in section Oligopterys to accommodate the finding that Aspicarpa, at least in part, is nested within Gaudichaudia. Three nothosections are named to accommodate new species described as amphiploids among sections. Gaudichaudia implexa is described as a new amphiploid species formed from lineages in section Trito- mopterys and section Gaudichaudia (nothosection Tritomochaudia). Gaudichaudia symplecta and Gau- dichaudia synoptera are described as two new amphiploid species formed from lineages in section Tri- tomopterys and section Cyclopterys (nothosection Cyclotomopterys). Gaudichaudia zygoptera and Gau- dichaudia intermixteca are described as two new amphiploid species formed from lineages in section Tritomopterys and section Zygopterys (nothosection Zygotomopterys). Gaudichaudia andersonii is de- scribed as a new amphiploid species formed from lineages within section Cycloptera. RESUMEN Se describen seis especies nuevas en el género Gaudichaudia y se revisa la taxonomia seccional en el género de acuerdo con los hallazgos del andlisis molecular del cloroplasto y de los genomas nucleares. Se guitan Gaudichaudia cycloptera, G. chasei y G. mcvaughii de la secci6n Gaudichaudia y se colocan en la secci6n Cyclopterys. Se quitan G. krusei y G. subverticillata de la secci6n Gaudichaudia y se colocan en la seccion Archaeopterys. Se pone Gaudichaudia hirtella, comb. nov. en la secci6n Oligopterys para acomodar el hallazgo de que Aspicarpa por lo menos en parte, se anida dentro de Gaudichaudia. Se nombran tres nothosecciones para acomodar la nuevas especies descrita como anfiploide entre sec- ciones. Se describe Gaudichaudia implexa como una nueva especie anfiploide formada de linajes en la seccion Tritomopterys y la secci6n Gaudichaudia (nothosecci6n Tritomochaudia). Se describen Gaudi- chaudia symplecta y Gaudichaudia synoptera como dos especies nuevas del anfiploide formada de linajes en la seccion Tritomopterys y la secci6n Cyclopterys (nothosecci6n Cyclotomopterys). Se describen Gau- dichaudia zygoptera y Gaudichaudia intermixteca como dos especies nuevas del anfiploide formada de linajes en la secci6n Tritomopterys y la secci6n Zygopterys (nothosecci6n Zygotomopterys). Se describe el Gaudichaudia andersonii como una especie nueva del anfiploide formado de linajes dentro de la seccién Cycloptera. Key Words: Amphiploid, Gaudichaudia, polyploid complex, Malpighiaceae, nothospecies, systematics. Gaudichaudia (Malpighiaceae) is a genus of woody vines, vining shrubs, and suffrutescent sub- shrubs inhabiting xeric to mesic habitats in Me- soamerica and northern parts of South America. Most of the diversity in Gaudichaudia is geograph- ically concentrated in central, western and southern Mexico with variable wide-ranging lineages reach- ing into northeastern and northwestern Mexico, and south into Central America and northern South America. Several regional and narrow endemics oc- cur in central Mexico south of the Tropic of Cancer and west of the Isthmus of Tehuantepec. Gaudi- chaudia was monographed by Franz Niedenzu in 1928, but little progress was made in understanding the genus until the work of Anderson in the 1980’s. Niedenzu attempted to coordinate the prior works of de Candolle, Adrien de Jussieu, Sessé and Mo- cino, Kunth, Chodat, Rose, and others who had de- scribed taxa in the genus in the previous century. Niedenzu himself added numerous names, con- structing an elaborate taxonomic system that in- cluded two subgenera and three sections covering eleven species and twenty-three infraspecific taxa. Although Niedenzu proliferated names in the ge- nus, he also significantly reduced the nomenclatural superfluity in Gaudichaudia by listing twenty-nine names from five genera in synonomy. While Nie- denzu’s work was a significant step in clarifying the systematics of Gaudichaudia, the profusion of names he introduced had the opposite effect. Nie- denzu’s taxa were largely based on narrow taxo- nomic concepts that relied on characters that are now clearly seen as variable within lineages and sometimes plastic even within a single individual. For example, he used the number of glands on the caylx to diagnose forms within varieties within sub- 238 species in both G. cycloptera and G. cynanchoides. Furthermore, Niedenzu’s reliance on floral features in Gaudichaudia, which like other Malpighiaceae is distinctive in the ancient (Taylor and Crepet 1987) and conservative features of its flowers (An- derson 1979), yielded a taxonomy that overlooked some distinctive species while emphasizing rela- tively minor variations in widespread taxa. To fur- ther confound taxonomic matters in Gaudichaudia, the herbarium specimens available to Niedenzu for his monograph were largely collected prior to 1910, and most of the collections he cited, even those designated as types, are lacking specific locality data. Some of the types are fragmentary or consist of immature stages, in a few cases without fruits or mature flowers. The names based on inadequate type material may never be clearly referable to a lineage and are therefore of little or no taxonomic value today. Taxonomy in Gaudichaudia has been revisited in recent years by Anderson (1987, 1993), who clar- ified some of the nomenclature and described three new species that were missed by Niedenzu and his predecessors. Anderson (1993) accepted Nieden- zu’s sectional taxonomy: section Tritomopterys, section Zygopterys, and section Gaudichaudia, all of which are distinguished on fruit wing morphol- ogy. Fruit wing morphology in section Tritomop- terys (G. albida Schlecht. & Cham., G. diandra (Nied.) Chodat, G. hexandra (Nied.) Chodat) is characterized by highly asymmetric lateral wings (Fig. 1), and in section Zygopterys (G. galeottiana (Nied.) Chodat) by distally rounded, free and sym- metric lateral wings, and a well developed posterior wing (Fig. 2). Section Gaudichaudia is character- ized by a rounded fruit wing with the apex some- times notched, but with lateral wings scarcely free at the apex and completely confluent at the base (Fig. 2). Anderson (1993) includes G. cynanchoides H. B. K., G. cycloptera (DC.) W. R. Anderson, G. subverticillata Rose, G. chasei W. R. Anderson, G. mcvaughii W. R. Anderson, and G. krusei W. R. Anderson in section Gaudichaudia (Fig. 2 in part, Figs. 3, 4). Anderson’s (1987, 1993) published work on Gaudichaudia recognizes ten species, in- cluding the three he described (G. mcvaughii, G. krusei, G. chasei). Anderson (1993) has shown that all ten of the recognized species are diploids with n = 40 meiotic chromosome pairs. Anderson (per- sonal communication) also recognizes several spe- cies in Gaudichaudia that have yet to be published. Three of those (species-A, “‘G. intermedia’; spe- cies-B, ““G hirsuta’; species-C, ““G. velutina’’) are clearly in section Tritomopterys and closely related to the G. albida—G. diandra—G. hexandra com- plex (Fig. 1). At least some of the undescribed spe- cies in section Tritomopterys are also known dip- loids as shown by Anderson’s (1993) chromosome counts published as G. albida sens. lat. Another undescribed species in Gaudichaudia is a very dis- tinctive close relative of G. cycloptera (species-D, MADRONO [Vol. 49 ““G. mexiae’’) that is narrowly restricted to lower elevations in remote areas on the west slopes of coastal Jalisco, Nayarit and Sinaloa (Fig. 3). Judg- ing from morphology of the samaras and the lim- ited chromosome counts available, all of these un- described elements of Gaudichaudia would be eas- ily accommodated within the sections established by Niedenzu as endorsed by Anderson. While the sectional taxonomy in Gaudichaudia based on fruit wing morphology is supported by overall morpho- logical similarity of the fruits, recent evidence from comparative analyses of the chloroplast and nuclear genomes (Jessup 1994, 2002), clearly shows that section Gaudichaudia as previously constructed is not monophyletic. In this paper I briefly summarize molecular evidence supporting sectional revisions in Gaudichaudia and propose a new combination resulting from the finding that Aspicarpa is, at least in part, nested within Gaudichaudia (Cameron et al. 2001; Davis et al. 2001). I propose three new sections and three new nothosections in Gaudi- chaudia and reclassify anomalous species. Six new species are described. MOLECULAR EVIDENCE Total genomic DNA was extracted from 118 Gaudichaudia specimens collected across a wide geographic region of Mexico. Extraction protocol and restriction site analysis followed procedures outlined in Dowling et al. (1996). Accession data and detailed laboratory procedure is presented in Jessup (1994). Restriction sites found on the chlo- roplast genome are presented in Table 1. Twenty- one informative sites were produced with 10 of the 20 enzymes screened. Thirty-four distinct chloro- plast haplotypes were discovered, each present in between 1 and 29 specimens. Restriction site data for representatives of each of the 34 haplotypes is given in Table 2. Phylogenetic analysis of the cpDNA data was performed with Hennig 86 v.1.5 (Farris 1989). Autapomorphies were excluded from the analysis. The procedure “‘mhennig*’’ was used to search for the shortest possible trees. Branch swapping on the shortest trees found by multiple initial passes with different OTU addition sequenc- es produced a set of trees that was then used as the starting point for procedure “‘bb’’, an extended branch-swapping algorithm that searches all possi- ble trees. As a check on thoroughness of the heu- ristic algorithms, a separate run of the procedure ‘ie; bb’’, an implicit enumeration algorithm fol- lowed by branch-swapping, was used to find all of the shortest trees, which were then used to compute the majority rule consensus tree using PAUP (Swofford 1990). The majority rule consensus tree of 26 equally parsimonious trees obtained from restriction site characters is presented in Fig. 5. Terminal branches in the cpDNA tree represent the 34 distinct ge- nomes discovered in the sample of 118 plants. Ro- 2002] JESSUP: GAUDICHAUDIA Section Tritomopterys @ G.albida A G.diandra 3+ G. hexandra (undescribed species) % G.sp. ined. “intermedia” ©. G.sp. ined. “hirsuta” ¥€ G.sp. ined. “velutina” Fic. 1. Geographic distribution of species and fruit morphology in representative collections from section Tritomop- terys. A-E, G. albida, A) Jessup 4041, Oaxaca, B) Jessup 4052, C) Anderson 13275, Mexico, D) Jessup 4067, Guerrero, E) Jessup 4056, Chiapas; F—J, G. diandra, F) Jessup 4088, Nayarit, G) Jessup 4066, Guerrero, H) Anderson 13309, Michoacan, I) Jessup 4032, Michoacan, J) Anderson 12937, Morelos; K—N, G. sp. ined. “‘intermedia’’, K) Anderson 13225, Chiapas, L) Jessup 4051, Oaxaca, M) Anderson 13224, Chiapas, N) Jessup 4055, Chiapas; O, G. sp. ined. “hirsuta”, Jessup 4048, Oaxaca; P-R, G. sp. ined. “‘velutina”’, P) Jessup 4058, Chiapas, Q) Jessup 4060, Chiapas, R) Jessup 4054, Oaxaca. Samara illustrations are approximately | x. 240 MADRONO [Vol. 49 Section Gaudichaudia @ G. cynanchoides {J OP - uty Zz Section Zygopterys Y G. galeottiana Fic. 2. Geographic distribution of species and fruit morphology in representative collections from section Gaudi- chaudia and section Zygopterys. A—L, section Gaudichaudia: A) Jessup 4078, Jalisco, B) Koch 82260, Mexico, C) Jessup 4077, Jalisco, D) Nicolas 5078, Puebla, E) Jessup 4075, Michoacan, F) Anderson 13296, Michoacan, G) Jessup 4111, Hidalgo, H) Anderson & Laskowski 3884, Michoacan, I) Jessup 4112, Hidalgo, J) Jessup 4109, Hidalgo, K) McVaugh 13204, Michoacan, L) Anderson & Laskowski, Michoacan; M-—Q, section Zygopterys: M—N) Galeotti X. 1844, Tutepeji, Puebla (type), O) Jessup 4038, Puebla, P) Anderson & Laskowski 4294, Puebla, Q) Anderson 13128, Oaxaca. Samara illustrations are approximately | x. bustness of the phylogenetic estimate is indicated by the fact that all but two branches are supported by 100% of the trees in the consensus calculation. Detailed geographic mapping of the cpDNA hap- lotypes and taxonomic assignment of plants carry- ing those chloroplast genomes is presented in Jes- sup (1994). The cpDNA phylogeny taken together with morphological features supports recognition of six sections in Gaudichaudia. Sections Tritomop- terys and Zygopterys as recognized by Anderson (1993) are well supported as monophyletic by the cpDNA analysis. Section Gaudichaudia is mono- phyletic with the removal of G. cycloptera, G. mevaughii, G. chasei, G. subverticillata, and G. krusei. The geographically widespread species G. cynanchoides is shown to be a genetically diverse assemblage of morphologically similar lineages. The methods used in this study were unable to re- solve the basal polytomy. The cpDNA phylogeny is illustrated as an unbranched network (Fig. 5) to emphasize the lack of phylogenetic resolution among the sections in Gaudichaudia. To accommodate the species removed from sec- tion Gaudichaudia two new sections are proposed here; section Cyclopterys and section Archaeopter- ys. Section Cyclopterys is well supported by the cpDNA analysis as monophyletic and includes G. cycloptera, G. mcvaughii, and G. chasei, as well as the undescribed elements discussed above. Section Archaeopterys is paraphyletic and, based on mor- phological evidence and evidence from analysis of nuclear DNA discussed below, comprises a cluster of phylogenetically basal lineages within the genus. There is not, however, sufficient evidence to root the cpDNA network with section Archaeopterys. Section Oligopterys is proposed here to accom- modate species of Aspicarpa that have been found to nest within Gaudichaudia, as reported by Cam- eron et al. (2001), Davis et al. (2001) and Jessup (1994). TAXONOMIC REVISIONS AND NEW SPECIES Revision of sectional taxonomy of the diploid species. Sectio Cyclopterys Jessup sect. nov. (Fig. 3)— TYPE: Mexico, Michoacan: Tuzantla, 110 km N Huetamo, rd to Zitacuaro, 670 m. 15 October 1988, Jessup 4033 (Holotype MICH; isotypes MO, UC, IEB). 2002] JESSUP: GAUDICHAUDIA 241 — - a Section Cyclopterys 66 G. cycloptera @ G. chasei OF G. mcevaughii G. sp. ined. “mexiae” G. mevaughii Fic. 3. Geographic distribution of species and fruit morphology in representative collections from section Cyclopterys. A-D, G. cycloptera, A) Jessup 4029, Guerrero, B) Jessup 4083, Jalisco, C) Jessup 4033, Michoacan, D) McVaugh & Koelz 534, Nayarit; E—G, G. chasei, E) Lyonnet 28, Morelos, F) Jessup 4009, Morelos, G) Bates 3426, Morelos: H) G. sp. ined. ““mexiae”, Croat 45249, Nayarit; I-J, G. mcvaughii, 1) Anderson 12699, Colima, J) Koch & Fryxell 82218, Guerrero. Samara illustrations are approximately 1X. Section Archaeopterys @ G subverticillata + G. krusei Fic. 4. Geographic distribution of species and fruit morphology in representative collections from section Archaeop- terys. A) G. subverticillata, Jessup 4087; B) G. krusei, Jessup 4069. Samara illustrations are approximately 1X. 242 TABLE 1. INFORMATIVE CPDNA RESTRICITON SITES FOUND IN 118 SPECIMENS. Each restriction site is represented by a letter code and referred to in the data matrix in Table 2. Enzyme Informative site(s) Site code Ban II 96> 82+ 1.4 Q Bcl I 4457+? H 3.6 > 2.1 + 1.5 O Se) => 20) SPL) T Cla I 19.0 > 8.2 + 10.8 E 2.8 > 1.2 + 1.6 F 2.95 1.3 + 1.6 G Dra I 6.15 > 5.48 + .67 K 5.48 > 4.2 + 1.28 L 31 Ss Io aP IS M Eco RI 3.5) => 340 =P OLS A Eco RV 16.5 > 9.5 + 7.0 D Hae III ALS <> 1.3 se 2? I Lab > 12 se OD J 16757?+? R Mgd) = EY S Hine II S19 => SD =P Dell P A232) => 2.3 a7 2O UW Hpa II 5.1 > 3.6 + 1.5 N Xba 3.8 > 2.0 + 1.8 B 30 => Zo) ar ILS (C To promote nomenclatural clarity, and to bring evidence from DNA analyses to bear on taxonomic circumscription of this section, under provisions of the ICBN Saint Louis Code, Article 22.6 (Greuter et al. 2000), I am designating a different holotype for section Cyclopterys than the type for the name on which the section is based, which is Gaudi- chaudia cycloptera (DC.) Anderson. The earliest name for the species according to Anderson (1987) is Hiraea? cycloptera DC. Prodr. 1: 586. 1824, and the type for that name is an unnumbered illustration among de Candolle’s collection of plates, referred to simply as “fl. mex. ic. ined.” (de Candolle 1824), a plate presumably copied from Sessé and Mocino’s original painting in the Jcones Florae Mexicanae, now at HU. The original watercolor il- lustration, the presumed virtual type, was titled, “Triopteris oblongifolia’ (HU accession number 6331.0888). Many of the 279 names based on types that are plates or copies of plates from the Sessé and Mocino expedition were never vouchered with a collection (McVaugh 1980). This is most likely the case for G. cycloptera. So far as I am aware there is no known specimen corresponding to the plate, though the possibility remains that a speci- men will be found in MA or among the collections belonging to a recipient of specimens collected on the Sessé and Mocino expedition. Establishment of section Cyclopterys is warranted on the evidence from DNA analyses together with morphological comparisons. I am designating a type for the sec- tion that can be subjected to further DNA analysis. The specimen designated as holotype for the sec- tion carries cpDNA haplotype Z, one of five hap- lotypes found in a well supported monophyletic G. MADRONO [Vol. 49 cycloptera (Fig. 5). G. cycloptera bearing cpDNA haplotype Z has not been discovered as a compo- nent in any amphiploid formed among sections in Gaudichaudia, although the closely related cpDNA haplotype AB is shared between some lineages within section Cyclopterys, in particular between G. cycloptera and G. mcvaughii, (Jessup 1994). Herbae fruticesve scandentes e basi fruticosa, caulibus longissimis volubilibus in vegetationem sustinentem extendentibus 4—12 m. Folia decussata et sensim decrescentia a basi ad ramos distales flo- rescentes, late elliptica ad ovata, subsessilia ad dis- tincte petiolata, basi hastata, cordata, truncata, vel cuneata. Indumentum partes maturas vegetativas to- tae plantae tegens, dense sericeum vel sparse hir- sutum ad grosse hispidum, e trichomatibus ramosis constans, trabeculis laevibus ad sinuosis, adpressis ad erecto-patentibus vel subaristatis. Flores omnes chasmogami, ad anthesin 1.5—3 cm lati, in umbellis geminatis 4-floris in axillis foliorum reductorum vel in dichasiis foliosis axillaribus dispositi, interdum in thyrsum elongatum distaliter fasciculati. Samarae orbiculares ad cordatae, 1—2 cm in diametro, alis lateralibus symmetricis, ad carpophorum confluen- tibus et rotundatis ad acutis vel late retusis, lobis apicalibus alarum obtusis ad acutis incisuram sinu lato ad angusto facientibus, vel alis lateralibus con- fluentibus ad apicem, margine distali itaque integra. Superficies dorsalis fructus alam distinctam inter- dum prominentem vel reticulum congestum pen- nularum supra nucem ferens. Vines and vining shrubs from a shrubby base, the longest twining stems reaching 4—12 m into the supporting vegetation. Leaves decussate and grad- ually reduced from the base to the distal flowering branches, broadly elliptic to ovate, subsessile to distinctly petiolate, leaf bases hastate, cordate, trun- cate or cuneate. Vesture thinly or densely sericeous or sparsely hirsute to coarsely hispid, comprising branched trichomes, the trabeculae smooth to sin- uose and appressed to erect spreading or subaris- tate, the indumentum covering mature vegetative parts of entire plant. Flowers all chasmogamous, 1.5—3 cm in diameter at anthesis, on paired four- flowered umbels in the axils of reduced leaves, or in axillary leafy dichasia, sometimes clustered dis- tally to form an elongate thyrse. Samaras orbicular to cordate, 1—2 cm in diameter, the lateral wings symmetric, confluent and rounded to acute or broadly retuse at the carpophore, the apical wing lobes obtuse to acute, forming a notch with a broad to narrow sinus, or the lateral wings apically con- fluent, the distal margin thus entire. The dorsal sur- face of the fruit bearing a distinct, sometimes prom- inent wing or a congested nexus of winglets over the nut. Phylogenetic analysis of restriction sites in the chloroplast genome of Gaudichaudia (Fig. 5) clear- ly supports Niedenzu’s section Tritomopterys con- taining lineages related to G. albida. The phylo- genetic analysis also supports section Gaudichau- 2002] JESSUP: GAUDICHAUDIA 243 TABLE 2. CPDNA RESTRICTION SITE MATRIX FOR HAPLOTYPES. Restriction site codes refer to Table |. Each haplotype is designated by 1-2 letters. Haplotype Specimen Estado A J4047 Oaxaca B J4059 Chiapas (C J4055 Chiapas D J4015 Morelos E J4036 Michoacan F J4056 Chiapas G J4103 Zacatecas H J4064 Guerrero I J4087 Nayarit JJ J4069 Guerrero K A13216 Oaxaca J J4027 Guanajuato M A3707 Jalisco N J4102 Sinaloa O J4008 Morelos P J4035 Michoacan Q J4037 Michoacan S J4081 Jalisco ie J4018 Morelos R J4007 Guanajuato U J4112 Hidalgo V J4042 Oaxaca W J4049 Oaxaca xX A12990 Oaxaca a’ J4029 Guerrero 7, J4033 Michoacan AA J4030 Michoacan AB J4024 Mexico AC A12699 Colima AD J4038 Puebla AE J4009 Morelos AF A4510 Guerrero AG J4039 Puebla Putative outgroup Apsicarpa Sites (refer to Table 1) ABCDEFGHIJKLMNOPORSTU 000101000000110000000 000111000000110000001 010111000000110000001 000111000100110000001 000111000100100000000 000110000100110000001 000111000000001010010 001111000000100000000 101111000000100000000 011111000000100000000 010111101000110100000 000111101000110100000 000111101000010100000 000110101000110100000 000111101000110000000 100111000000101011011 OTA ARG OOOO CAs AO mea Os A AO) IE OONONCNOMONONGMENO)AL SE O}AEO) TL) LNAI ITO O)ONCNONON ONC) ALTOHAL 110) 4116) LOLTEALOOCCOPOOLOLLOT0 101111000000001011010 001111010010100000100 101111010010100000100 101111000010100000100 101001000000100000000 101001000000100000000 001001000000100000000 101011000000100000000 101010000000100000000 101111000001100000101 101111000000100000000 101111000000000000000 001111000001100000101 101111000000100000000 dia containing lineages related to G. cynanchoides, as well as section Zygopterys containing the single species, G. galeottiana. In the phylogenetic analy- sis section Zygopterys and section Tritomopterys were each found to be monophyletic. Based on the molecular data, however, a monophyletic section Gaudichaudia cannot include G. cycloptera, G. mevaughii, G. subverticillata, G. krusei, or G. chas- ei as proposed by Anderson (1993). Those species are therefore moved to new sections as detailed be- low. Section Cyclopterys is strongly supported as monophyletic by the chloroplast DNA phylogeny, comprising the distinctive and geographically wide- spread type species, G. cycloptera and the narrow endemic, G. mcvaughii, known only from scattered and narrowly delimited low elevation localities on the Pacific coast of Colima, Guerrero, and Oaxaca. In addition to sharing, in some populations, a dis- tinct chloroplast haplotype with G. cycloptera, G. mcvaughii also shares morphological features of the fruit and habitat attributes with G. cycloptera. The elaboration of a dorsal wing in the fruits of both species, and the size and shape of the fruits provide further evidence that these taxa are closely related. Both G. cycloptera and G. mcvaughii are plants of mesic understory or forest edge thickets. Both are known diploids without cleistogamous flowers. Although the branch supporting the single col- lection of G. chasei included in the analysis (Fig. 5) is unresolved with respect to branches support- ing sections, G. chasei has several morphological features in common with members of section Cy- clopterys, notably the expanded dorsal keel on the samara, and lateral fruit wings that are basally con- fluent. Gaudichaudia chasei, which Anderson (1987) argues is morphologically close to G. mevaughii, is known from only a small area in Mo- relos. The habitat where it grows is more similar to the mesic understory habitat of G. cycloptera from adjacent Edo. Mexico than the habitat of the typical species of section Gaudichaudia (G. cynanchoi- des), which grows in the xeric matorral of the Al- 244 MADRONO [Vol. 49 aie 13 ay) ; mexia ARCHAEOPTERYS cycloptera mevaughii subverticillata Oo) @) @ AB) CO chasei CYCLOPTERYS OLIGOPTERYS (AF) cynanchoides sens. lat. hirtella galeottiana Majority rule consensus phylogeny of cpDNA haplotypes in Gaudichaudia. Fic. 5. Majority rule consensus phylogeny of cpDNA haplotypes showing the relationship of sections in an unrooted network. Each terminal branch represents a distinct cp DNA haplotype defined by a unique configuration of restriction sites. Letters in circles designate haplotypes referred to in Table 2. Representative fruits of species of non-hybrid origin illustrate the range of fruit morphology in each section. Position of the fruits relative to branches of the network is approximate. toplano Mexicano (Fig. 2). Gaudichaudia chasei, like G. cycloptera and G. mvaughii, lacks the cleis- togamous flowers that are a prevalent feature in G. cynanchoides. Though perhaps a divergent member of section Cyclopterys, G. chasei clearly fits better here than in the other sections and is not otherwise sufficiently distinct to justify a separate section. The molecular evidence, absence of cleistogamy and other morphological evidence mentioned by Anderson (1987), the similarity of geographic range and habitats, and the evidence from Ander- son’s (1993) study of chromosomes, indicating that all are diploids with n = 40 meiotic chromosome pairs, taken together strongly supports recognition of section Cyclopterys, containing G. cycloptera, G. mevaughii, G. chasei, and the undescribed lineage from Nayarit and Jalisco (G. sp. ined. “‘mexiae’’) as a discernable monophyletic clade within Gau- dichaudia. Section Gaudichaudia sensu Anderson (1993) included G. subverticillata and G. krusei, two spe- cies that he argued are closely related (Fig. 4). Both are subshrubs with rounded fruit wings resembling in outline the fruits of G. cynanchoides. The phy- logeny based on chloroplast genomes does not, however, support inclusion of those species in a monophyletic section Gaudichaudia. The fruits are generally similar in size and shape to those of sec- tion Gaudichaudia, and are quite distinct from fruits of species included in section Tritomopterys. Nevertheless, G. subverticillata and G. krusei are unambiguously placed as outgroups to the G. al- 2002] bida complex (section Tritomopterys) in the parsi- mony analysis of the chloroplast genomes (Fig. 5). Taken together, G. subverticillata, G. krusei and section Tritomopterys are monophyletic. In view of the substantial divergence in gross morphology be- tween G. subverticillata and G. krusei on the one hand and members of section Tritomopterys on the other, however, their assignment to section Trito- mopterys is untenable. Gaudichaudia subverticil- lata and G. krusei are certainly similar in mor- phology: both are suffrutescent shrublets lacking cleistogamy, and both are known diploids. They oc- cupy similar habitats, and are both narrow endem- ics in a region of southwestern Mexico rich in nar- row endemics from many groups of plants, a fact that suggests the area may have served as an an- cient refugium where phylogenetically basal line- ages might be expected to persist. Furthermore, the comparative analysis of nuclear genomes based on randomly amplified DNA (Jessup 2002) places G. subverticillata and G. krusei as sister taxa close to members of section Cyclopterys, but on the periph- ery of the minimum spanning tree of Jaccard dis- tances. The evidence, taken together, corroborates the hypothesis that G. subverticillata and G. krusei are basal within Gaudichaudia. They are apparently closely related to each other, and more closely re- lated to section Tritomopterys and section Cyclop- terys than they are to section Gaudichaudia. To ac- commodate sectional placement of G. subverticil- lata and G. krusei the following new section is es- tablished. Sectio Archaeopterys S. L. Jessup sect. nov. Fruticuli suffrutescentes raro ramosi, ramis max- imis plerumque 0.5—1.0 m altis e basi lignosa. Folia brevipetiolata ad subsessilia, binata vel verticillata terna. Indumentum partes maturas vegetativas totae plantae tegens, sparse vel dense sericeum ad velu- tinum, e trichomatibus ramosis constans, trabeculis laevibus ad sinuosis et adpressis ad erecto-patenti- bus. Flores omnes chasmogami, in umbellis 4-floris verticillatis e nodis distalibus caulium primorum vel in dichasiis brevibus axillaribus a foliis caulinis superioribus subtentis dispositi. Samarae orbicular- es ad cordatae, alis lateralibus symmetricis, proxi- maliter acutis vel ad carpophorum infirme retusis, lobis distalibus alarum obtusis ad acutis, incisura apicali sinum obtusum ad acutum facienti, superfi- cies dorsalis fructus alam rudimentariam ad prom- inentem vel reticulum humile pennularum supra nu- cem ferens. Suffrutescent seldom branching shublets, the largest branches mostly 0.5—1.0 meter high from a woody base. Leaves short petiolate to subsessile, paired or in whorls of three. Vesture thinly or densely sericeous to coarsely velutinous, compris- ing branched trichomes, the trabeculae smooth to sinuose and appressed to erect spreading covering mature vegetative parts of entire plant. Flowers all JESSUP: GAUDICHAUDIA 245 chasmogamous, in verticillate four-flowered umbels from distal nodes of main stems, or on short axil- lary dichasia subtended by the upper stem leaves. Samaras orbicular to cordate, the lateral wings sym- metric, proximally acute to weakly retuse at the car- pophore, the distal wing lobes obtuse to acute, the apical notch forming an obtuse to acute sinus, the dorsal surface of the fruit bearing a rudimentary to prominent wing or low nexus of winglets over the nut. Type: Gaudichaudia subverticillata Rose The problem of long branch attraction makes in- terpretation of branching order among sections in Gaudichaudia problematic. Basal lineages with few close relatives in a genus of otherwise closely re- lated species complexes are particularly sensitive to inaccurate placement. Placement of section Ar- chaeopterys should therefore be tempered with skepticism until Tribe Gaudichaudieae can be ana- lyzed as a whole with sequence data chosen spe- cifically for this problem. With the foregoing sectional reassignments, sec- tion Gaudichaudia now comprises only vines with both chasmogamous and cleistogamous flowers producing rounded, essentially symmetric (cynan- choid) samaras. Section Gaudichaudia as revised includes only G. cynanchoides sens. lat., which is widespread in mesic to xeric ruderal habitats on the Altoplano Mexicano (Fig. 2). Although the only ac- cepted species remaining in section Gaudichaudia is G. cynanchoides, that name as now used encom- passes a diverse assemblage of microspecies prop- agating largely through abundant production of cleistogamous fruits. As evident in the series of dis- tinct chloroplast haplotypes and diverse morpho- types (Jessup 1994), G. cynanchoides is a species complex that should eventually be resolved into several closely related but geographically distinct species or subspecies. In addition to supporting the realignment of sec- tional taxonomy of recognized members of Gau- dichaudia, the results of molecular research clearly indicate that the genus Gaudichaudia itself, as con- ventionally delimited, is paraphyletic. My molecu- lar studies (Jessup 1994, 2002) included the genus Aspicarpa as the outgroup in several analyses, but in the maximum likelihood analysis (unpublished results) Aspicarpa was found to nest within Gau- dichaudia in a polytomy with section Cyclopterys, section Archaeopterys, and section Gaudichaudia. That result is corroborated by recent molecular phylogenies of the Malpighiaceae (Cameron et al. 2001; Davis et al. 2001) showing that at least some species now included within Aspicarpa are nested within Gaudichaudia. To accommodate the evi- dence demonstrating that Aspicarpa is, at least in part, nested within Gaudichaudia, | here establish a new section in Gaudichaudia to include those el- ements of Aspicarpa that are properly considered 246 species within a monophyletic Gaudichaudia. The type of the new section is established by the fol- lowing new combination. Gaudichaudia hirtella (Rich.) Jessup, comb. nov. Aspicarpa hirtella Rich., Mem. Mus. Paris 2: BID, Nest). Aspicarpa urens Lagasca, Gen. Sp. Pl. Nov. 1. 1816. Aspicarpa pruriens Desv., Desf. Cat. Hort. Paris. Gl, Bo ZIBB, M329). Gaudichaudia urens Chodat in Bull. Soc. Bot. Geneve Z Sew IDX, IOI, Section Oligopterys Jessup sect. nov. Fruticuli suffrutescentes caulibus paucis ad mul- tis erectis vel decumbentibus e basi lignosa. Folia opposita vel verticillata, basi rotundata ad cordata, sessilia ad subsessilia. Flores et chasmogami et cleistogami. Fructus sine ala laterali, nuculum oblique affixum sine carpophoro producentes, crista vel jugo dorsali instructi, sine ala dorsali. Type: Gaudichaudia hirtella (Rich.) Jessup, comb. nov. Suffrutescent shrublets with few to many erect to decumbent or trailing stems from a woody base. Leaves opposite or whorled, rounded to cordate at base, sessile to subsessile. Flowers both chasmo- gamous and cleistogamous. Fruits lacking a lateral wing, forming an obliquely attached nutlet lacking a carpophore, with a low dorsal crest or ridge, with- out a dorsal wing. Other species now placed in Aspicarpa will like- ly emerge as elements of Gaudichaudia. Only G. hirtella is included here in section Oligopterys since that was the species included in my study of chloroplast DNA phylogeny in Gaudichaudia. Al- though the chloroplast DNA parsimony analysis majority rule consensus tree does not resolve the branch supporting Aspicarpa with respect to branches supporting other sections, Aspicarpa is distinct enough in fruit morphology and plant habit to warrant a separate section. Circumstantial evi- dence from studies with labeled RAPD probes (Jes- sup 2002) support placement of section Oligopterys close to section Zygopterys. When amplified PCR products from G. galeottiana were probed against blots of RAPD gels in that study, lanes representing Aspicarpa hybridized the probe along with mem- bers of section Zygopterys and the intersectional amphiploids involving section Zygopterys. The probe also weakly hybridized the lane representing G. krusei, but did not hybridize lanes representing other lineages within Gaudichaudia. In the phenetic analysis of randomly amplified DNA (Jessup 1994) Aspicarpa clustered with members of section 77i- tomopterys on the minimum spanning tree of Jac- card distances. Recent DNA sequence analysis of generic phylogeny in Malpighiaceae (Cameron et MADRONO [Vol. 49 al. 2001; Davis et al. 2001) found Aspicarpa nested with members of section Tritomopterys. Definitive resolution of the phylogentic placement of sections in Gaudichaudia must, however, be deferred until DNA sequences for a broader sample of represen- tative taxa in tribe Gaudichaudieae are available. Taxonomic revisions in the amphiploid complex- es. Anderson began the task of resolving reticulate ancestry in Gaudichaudia. In addition to his de- scriptions of new species and chromosome counts in Gaudichaudia, Anderson explored the relation- ship of Gaudichaudia to other genera in Tribe Gau- dichaudieae (Anderson 1985) and contributed to an understanding of the genus through studies of re- productive life history traits (Anderson 1980). In particular, cryptic self fertilization, which occurs in distinctive cleistogamous flowers, is common to all lineages examined so far that have n = 80 meiotic chromosome pairs. All of the plants examined thus far that bear fruits with morphologies intermediate between those of plants fitting neatly into the sec- tions of Gaudichaudia (as herein defined) are tet- raploids, and they all bear cleistogamous flowers. The evidence presented by Anderson strongly sup- ports his hypothesis that plants bearing fruits with intermediate morphologies are amphiploids or the products of amphiploids formed among lineages in different sections of Gaudichaudia. Results sup- porting that hypothesis have been corroborated by the results of molecular studies (Jessup 1994, 2002). The amphiploids present a bewildering mélange of morphological variation that has heretofore been reticent to clean cut species delimitations. The com- bination of evidence now available from molecular studies and a morphological survey of a large num- ber of collections from across the geographic dis- tribution of Gaudichaudia in Mexico, however, re- veals several discrete elements among the tetra- ploids that can be clearly discerned and described as new species in Gaudichaudia. Five of the spe- cies described here are amphiploids formed among lineages from different sections, and one is appar- ently formed as an amphiploid among lineages within a section. In each case, the amphiploid ori- gin of the new species is supported by evidence from molecular studies. These are not simple F, hybrids, but rather wide-ranging lineages that prop- agate via selfing through cryptic self fertilization while maintaining outcrossing through chasmoga- mous flowers. Floral morphology is remarkably uniform across the Tribe Gaudichaudieae, and clearly fits the family-specific floral syndrome as- sociated with oil bee pollination (Buchmann 1987; Vogel 1990). Undescribed diploid lineages identi- fied by Anderson (Figs. 1, 3) are introduced and discussed in this paper in connection with their roles in formation of amphiploids among lineages in different sections of Gaudichaudia, and in delim- iting new taxa with which they might be confused. 2002] Only amphiploid lineages with unambiguous mor- phological attributes and clear support from the molecular research are described in this paper. Sev- eral additional amphiploid species not treated here remain undescribed in Gaudichaudia. Nothosections and new amphiploid species. Pro- visions are made in the ICBN (Greuter et al. 2000) for naming nothotaxa, taxa of known hybrid origin, and those provisions are codified in the St. Louis Code, Appendix I. A diagnosis or description is not required for the naming of notho-subdivisions of genera, and such names do not have types, but the ICBN requires that names of the parental taxa are specified when the name of the nothotaxon is pub- lished (Article H.9). However, species that are known or suspected to be of hybrid origin need not be designated as nothospecies and may be desig- nated as species (Article H.3). Nothosections are proposed here to contain Gaudichaudia lineages that are clearly the products of intersectional hy- bridization. The combined evidence from morphol- ogy, chromosomal counts and molecular studies supports designation of nothosections with unam- biguous specification of the sectional source of pa- rental lines, even to the point of specifying section- al contributions of the pollen and ovules for indi- vidual members of species within the nothosec- tions. Sectional sources of pollen and ovule parents are indicated below for the types and paratypes where known. It has not, however, yet been feasible to unambiguously identify species-level lineages contributing to formation of the species in the no- thosections. The species named below can be clear- ly circumscribed and assigned to nothosections, but which of several possible combinations of species within the parental sections gave rise to them re- mains unknown or ambiguous. In some cases the member species in nothosections proposed here are likely the products of complex interbreeding among several independently evolving lineages from each of the contributing parental sections. With those reasons in mind the species proposed here are named as species rather than nothospecies. Nothosection Tritomochaudia S. L. Jessup notho- sect. nov. (Gaudichaudia sect. Gaudichaudia X Gaudichaudia sect. Tritomopterys), Fig. 6. Pres- ently I am recognizing only a single broadly de- limited species. The molecular data, the chro- mosome data, and the geographic distribution of morphological variation suggest that this taxon comprises a swarm of autogamously propagating amphiploid microspecies that retain viable chas- mogamy. Gaudichaudia implexa S. L. Jessup, sp. nov. (Fig. 6)—TYPE: Mexico. Jalisco: south shore of Lago de Chapala, 6.4 km W of Jalisco/Michoacan state line, 1620 m, dry thorn scrub in hills above lake, 3 Nov 1988, Jessup 4076 (Holotype MICH; iso- types CHAP, IEB, UC). JESSUP: GAUDICHAUDIA 247 Haec stirps variabilissima a Gaudichaudia cy- nanchoides secedit ala samarae lobos laterales an- ticos fere symmetricos ad valde asymmetricos acu- tos ad obtusos rotundatosve formanti, lobis sinu ad basin acuto ad obtuse angulato separatis, sinu in- terdum denticulo e margine antica alae lateralis vel in basin sinus ex apice nucis orienti, denticulo raro in lobum apicalem rotundatum inter lobos laterales alae crescenti. Samara autem Gaudichaudia cynan- choides ala laterali margine apicali fere integra gau- det, et quamquam ala incisuram apicalem parvam exhibet, samara ejus cynanchoidea est et lobis dis- cretis lateralibus sinu profundo separatis numquam instruit. Gaudichaudia albida, Gaudichaudia dian- dra et stirpes affines sectionis Tritomopterygos stat- im distinguendae sunt lobis lateralibus alae samarae valde asymmetricis, et cauda angusta postica alae Samarae vix vel haud confluenti cum ala laterali. Haec species interdum valde simulabit Gaudichau- dia galeottiana vel Gaudichaudia zygoptera aut alis lateralibus symmetricis rectis aut lobo lato postico constricto ad nucem aut utroque, sed clare distin- guenda est limbo texturae inter alas laterales al- amque posticam et alis lateralibus apicibus obtuse acutis pro alis plerumque latioribus rotundiorib- usque Gaudichaudia galeottiana et Gaudichaudia zygoptera; ubicumque autem admiscet speciebus il- lis, hae differentiae vix discernendae erit. Chro- mosomatum numerus, n = 80. This highly variable species is separable from G. cynanchoides by the samara wing forming two ap- proximately symmetric to strongly asymmetric acute to obtuse or rounded anterior lateral lobes apically separated by a sinus that varies from acute to obtusely angular at the base, the sinus sometimes with a small tooth originating from the antical mar- gin of the lateral wing or in the base of the sinus from the crown of the nut, the tooth rarely devel- oping as a rounded apical lobe between the lateral lobes of the wing. By contrast the samara in G. cynanchoides has the apical margin of the lateral wing largely entire, and though it occasionally has a small apical notch in the samara wing, the samara is cynanchoid and never has discrete lateral lobes separated by a deep sinus. Gaudichaudia albida, G. diandra and related species in section Tritomopter- ys can be immediately distinguished by the strongly asymmetric lateral lobes of the samara wing, and by the narrow posterior tail of the samara wing that is scarcely or not at all confluent with the lateral wing. In some cases this species closely resembles G. galeottiana or G. zygoptera in the development of symmetric upright lateral samara wings or a broad postical lobe constricted where it attaches to the nut. A few specimens bear both of those char- acteristics, but it is clearly distinguished by a flange of tissue between the lateral wings and the postical wing, and by the lateral wings with bluntly pointed apices rather than the generally broader more rounded wings of G. galeottiana and G. zygoptera. However, where it hybridizes with those species the 248 Fic. 6. MADRONO [Vol. 49 Nothosection Tritomochaudia G. implexa Gaudichaudia implexa fruit morphology and geographic distribution of the holotype and paratypes. A—K, plants with 2 ancestor from section Gaudichaudia: A) Jessup 4076 (type), B) Rzedowski 32522, C) Jessup 4079, D) Jessup 4080, E) Jessup 4081, F) Jessup 4082, G) Jessup 4037, H) Jessup 4018, 1) Jessup 4100, J) Anderson 12624, K) Jessup 4105; L—X, plants with 2 ancestor from section Tritomopterys: L) Jessup 4006, M) Jessup 4108, N) Anderson & Laskowski 3707, O) Jessup 4115, P) Anderson & Laskowski 4293, Q) Anderson & Laskowski 4056, R) Jessup 4002, S) Anderson 13316, T) Jessup 4000, U) Jessup 4001, V) Jessup 4113, W) Jessup 4114, X) Jessup 4106. Samara illustrations are approximately 1X. distinctions will be difficult to discern. I collected flower buds of the type, prepared aceto-carmine squashes of meiotic pollen mother cells and count- ed the chromosomes. In three separate preparations I found n = 80 pairs. Anderson (1993) reported chromosome counts in several collections included here as paratypes. Anderson found meiotic chro- mosome number was n = 80 in Rzedowski 32522, Anderson & Laskowski 3707, Anderson & Las- kowski 4056, Anderson & Laskowski 4293, Ander- son 12624, and Anderson 13316. Paratypes. Mexico. Plants with 2 ancestor from section Gaudichaudia: Hidalgo: 7 km NE Mezqui- titlan, rd to Zacualtipan, 1800 m, 17 Nov 1974, Rzedowski 32522 (MICH, IEB). Jalisco: Cerro Vie- jo, trail S of Tlajomulco, 1800 m, 5 Nov 1988, Jessup 4079 (CHAP, IEB, MICH, UC); 10 mi NE of Cocula, between Guadalajara and Autlan, 1380 m, 6 Nov 1988, Jessup 4080 (CHAP, IEB, MICH, UC); 10 mi NE of Cocula, between Guadalajara and Autlan, 1380 m, 6 Nov 1988, Jessup 4081 (CHAP, IEB, MICH, UC); 11.3 km NE of Tecol- otlan, between Guadalajara and Autlan, 1480 m, 6 Nov 1988, Jessup 4082 (CHAP, IEB, MICH, UC). Michoacan: Mpio. Benito Juarez, 17.5 km S of Zi- tacuaro, 0.5 km S of Guanoro, 500 m, 16 Oct 1988, Jessup 4037 (CHAP, IEB, MICH, UC). Morelos: 1 km SE of Laureles village limit, Barranca Tezahu- 2002] ate, ca. 3 km NW of Tlayacapan, 1800 m, 6 Oct 1988, Jessup 4018 (CHAP, IEB, MICH, UC). So- nora: 18 km N of Yécora—Hermosillo hwy, on rd to Sahuaripa, 850 m, 12 Sep 1990, Jessup 4100 (CHAP IEB. MICH, UC)...Zacatecas: 21 km_S. of Villanueva on rd to Jalpa, 1900 m, 12 Sep 1983, Anderson 12624 (MICH): 21 km S of Villa Nueva, 1900 m, 16 Sep 1990, Jessup 4105 (CHAP, IEB, MICH, UC). Plants with 2 ancestor from section Tritomopterys: Guanajuato: 16 km NE of San Fi- lipe, 1890 m, 2 Oct 1988, Jessup 4006 (CHAP, IEB, MICH, UC); 19 km E of San Luis de la Paz on rd to Victoria, 2070 m, 17 Sep 1990, Jessup 4108 (CHAP, IEB, MICH, UC). Jalisco: 9.7 km E of Vil- la Corona, above Lago Atotonilco, 1420 m, 24 Sep- tember 1966, Anderson & Laskowski 3707 (MICH). Nuevo Leon: 6.1 km S of Allende, between Linares and Monterrey, 10 Oct 1990, Jessup 4115 (CHAP, IEB, MICH, UC). Oaxaca: 7.4 km NE of Chazum- ba, 2150 m, 23 Nov 1966, Anderson & Laskowski 4293 (MICH). San Luis Potosi: 32.9 km W of Cd Valles, 485 m, 18 Oct 1966, Anderson & Laskowski 4056 (MICH); 3.2 km W of Cuidad Valles, 420 m, 1 Oct 1988, Jessup 4002 (CHAP, IEB, MICH, UC); 20 km E of Santa Catarina, 1200 m, 29 Oct 1983, Anderson 13316 (MICH). Tamaulipas: hwy 101, 1.6 km S of bridge over Rio San Marcos, 14.5 km S of ject with hwy 85, 30 Sep 1988, Jessup 4000 (CHAP, IEB, MICH, UC); 6.4 km N of jct hwy 85 and hwy 40, N of Guayalejo, 750 m, 1 Oct 1988, Jessup 4001 (CHAP, IEB, MICH, UC); NW of Tampico, 4.8 km E of Gonzalez, 8 Oct 1990, Jessup 4113 (CHAP, IEB, MICH, UC); 77 km N of Ciudad Victoria, 32 km N Rio Purificacion, 250 m, 8 Oct 1990, Jessup 4114 (CHAP, IEB, MICH, UC). Za- catecas: Mpio. Jalpa, 5.1 km E of jct hwy 54 and hwy 70, rd to Aguascalientes, 16 Sep 1990, Jessup 4106 (CHAP, IEB, MICH, UC). Gaudichaudia implexa is morphologically more variable and geographically more widespread than any other species in Gaudichaudia. The name, which means “‘entangled,”’ refers not only to the typical habit of the plant in relation to the surround- ing supportive, often spiny, prickly or thorny veg- etation, but to the fact that lineages within this spe- cies represent the entangled accretion of genetically intertwining amphiploid lineages formed among diploid species in the G. albida sens. lat. and the G. cynanchoides sens. lat. complexes. All of the lineages in this species proliferate via autogamous- ly produced samaras, exhibit samara wing mor- phology with some degree of shape intergradation among shapes typically found in the diploids, and on analysis of cpDNA exhibit a chloroplast haplo- type typical of either the G. albida species complex or the G. cynanchoides species complex (Jessup 1994, 2002). Judging from the spectrum of samara wing morphology, the broad range of stem and leaf vesture, and the combination of cpDNA haplotypes found it seems likely that more than one of the species in section Tritomopterys (Fig. 1) is active JESSUP: GAUDICHAUDIA 249 in formation of the amphiploids. Ten cpDNA hap- lotypes were detected in samples representing sec- tion Tritomopterys, and twelve of the G. implexa collections (paratypes) shared one of those haplo- types with three species in section Tritomopterys (G. albida, G. diandra, Gaudichaudia sp. ined. *‘in- termedia’’). One additional haplotype closely relat- ed to other haplotypes carried by species in section Tritomopterys (Gaudichaudia sp. ined. “‘velutina,”’ Gaudichaudia sp. ined. “‘intermedia,”’ and G. dian- dra) was detected in a single collection of G. im- plexa. Of the eight cpDNA haplotypes detected in samples representing section Gaudichaudia, eight of the G. implexa collections shared one of those haplotypes with G. cynanchoides. Three additional haplotypes carried by specimens representing sec- tion Gaudichaudia were each detected in one col- lection of G. implexa. The plants are typically found in ruderal habitats and range in geographic distribution (Fig. 6) from southern Puebla westward through the Eje Volcan- ico Transversal to central Jalisco, northward on the Altiplano Mexicano to Nuevo Leon, in scattered locations in the Sierra Madre Occidental, and east- ward into the Sierra Madre Oriental in Tamaulipas, San Luis Potosi, Queretaro, and Hidalgo. Anoma- lous collections from Chihuahua, Coahuila and Du- rango probably represent rare isolated lineages of this species, similar to the northern disjunct popu- lation sampled from southeastern Sonora. Though formed as amphiploids among lineages in section Tritomopterys and section Gaudichaudia, the geo- graphic range and apparent ecological amplitude of G. implexa far exceeds that of either ancestral dip- loid lineage. Nothosection Zygotomopterys S. L. Jessup notho- sect. nov. (Gaudichaudia sect. Tritomopterys X Gaudichaudia sect. Zygopterys), Fig. 7. This no- thosection contains two amphiploid species from southern Puebla and west-central Oaxaca, with an isolated population of one species from cen- tral Guerrero. Gaudichaudia zygoptera S. L. Jessup, sp. nov. (Fig. 7)—TYPE: Mexico. Oaxaca: 10.1 km N of hwy 190 on rd to Guelatao, 1880 m, in thicket near stream, 22 Oct 1988, Jessup 4042 (Holotype MICH; isotypes CHAP, IEB, UC). A Gaudichaudia galeottiana facile distinguenda lobis lateralibus alae samarae valde asymmetricis, uno vel utroque lobo majori [quam eis Gaudichau- dia galeottiana], lobo postico angustiori, et nulla constrictione lobi postici ad nucem, vel aliquot his differentiis. A formis Gaudichaudia implexa lobum posticum alae samarae latum ferentibus distinguen- da apicibus loborum lateralium alae samarae late rotundatis pro apicibus obtuse acutis Gaudichaudia implexa. Readily distinguished from G. galeottiana by a marked asymmetry in the lateral lobes of the sa- 250 Fic. 7. MADRONO [Vol. 49 Nothosection Zygotomopterys © G. intermixteca wy SB e® a | 3+ G. zygoptera Nothosection Zygotomopterys fruit morphology and geographic distribution of the holotypes and paratypes. A-E G. intermixteca: A) Jessup 4040, B) Anderson 13031, C) Jessup 4043, D) Jessup 4047 (type), E) Jessup 4046, F) Jessup 4045; g—m, G. zygoptera: G) Anderson 12990, H) Jessup 4044, 1) Jessup 4038, J) Jessup 4072, K) Jessup 4042 (type), L) Jessup 4049, M) Anderson 13138. Samara illustrations are approximately 1x. mara wing, by one or both lobes larger than those found in G. galeottiana, by a narrower postical lobe of the samara wing, and by the absence of a prom- inent constriction in the postical lobe where it at- taches to the nut, or by some combination of these features. Distinguished from phases of G. implexa that bear a broadened postical lobe of the samara wing by the broadly rounded apices on the lateral lobes of the samara wing, in contrast with the gen- erally bluntly acute apices typical of the lateral lobes in G. implexa. Paratypes. Mexico. Guerrero: 103 km N of Ac- apulco on rd to Chilpancingo, 8 km along rd to El Alquitran, 2000 m, 30 Oct 1988, Jessup 4072 (CHAP, IEB, MICH, UC). Oaxaca: 22.5 km S of Huahuapan on rd to Oaxaca, 2100 m, 20 Oct 1988, Jessup 4049 (CHAP, IEB, MICH, UC); Mpio. Oa- xaca, vicinity of Monte Alban, 21 Oct 1988, Jessup 4044 (CHAP, IEB, MICH, UC). Puebla: .8 km NW of Cacaloapan on rd between Puebla and Tehuacan, 1970 m, 19 Oct 1988, Jessup 4038 (CHAP, IEB, MICH, UC); 89 km S of Teotitlan on rd to Oaxaca, 1500 m, 10 Oct 1983, Anderson 12990 (MICH); 3 km S of Ocotlan on rd from Oaxaca to Puerto An- gel, 1540 m, 15 Oct 1983, Anderson 13138 (MICH). Plants of ruderal habitat ranging from central and southern Puebla south into central Oaxaca (Fig. 7). The name refers to the zygomorphy evident in the lobes of the lateral samara wing, which indicates the influence of ancestral hybridization with line- ages from section Tritomopterys. Though the evi- dence is weak for the branching order of sections, in the maximum likelihood analysis of cpDNA re- striction sites section Zygopterys and section Tri- tomopterys form a monophyletic group with section Tritomopterys haplotypes forming a crown cluster, indicating possible paraphyly of section Zygopter- ys. All but one of the specimens of G. zygopterys sampled for cpDNA haplotypes carry haplotypes closely related to that carried by G. galeottiana, indicating that most of the hybridization contribut- ing to the formation of this species involves pollen donors from section Tritomopterys. The paratype Anderson 12990 has n = 80 chromosome pairs at meiosis (Anderson 1993). Only one of the cpDNA haplotypes encompassed in section Zygopterys is unequivocally assigned to G. galeottiana. The other haplotypes were found exclusively in G. zygoptera. The possibility re- mains that variation seen in samara morphology in G. zygoptera is indicative of relictualism, repre- senting a tendency toward increased zygomorphy in the samara wing that persists in lineages that were ancestral to section Tritomopterys. It seems more probable, however, that cpDNA haplotype di- 2002] versity in section Zygopterys is indicative of ances- tral diversity within G. galeottiana that has been swamped by introgression with adventive lineages from section Tritomopterys. One population of G. zygoptera, the western disjunct in central Guerrero, carries a cpDNA haplotype shared with three line- ages (G. albida, G. diandra, Gaudichaudia sp. ined. “‘intermedia’’) within section Tritomopterys. An anomalous plant collected in southern San Luis Potosi (Anderson & Laskowski 4043) that is mor- phologically well placed in G. implexa bears a cpDNA haplotype from section Zygopterys. That plant, growing 500 km north of the range of G. zygoptera, is probably a hybrid between G. zyg- optera and either G. cynanchoides or G. implexa. The nuclear DNA signature, based on randomly amplified DNA, places it close to G. implexa, nest- ed close to G. cynanchoides (Jessup 1994). Gaudichaudia intermixteca S. L. Jessup, sp. nov. (Fig. 7)—TYPE: Mexico. Oaxaca: Mpio. Oaxa- ca, vicinity of Monte Alban, 21 October 1988, Jessup 4047 (Holotype MICH; isotypes CHAP, IEB, UC). A stirpibus nothosectionis Cyclotomopterys, qui- buscum generatim similis est forma alae samarae, distinguenda est lobo postico angustiori, apice lobi postici latiori minusque acuto-acuminata, lobis la- teralibus vix ad valde asymmetricis, et samaris la- tioribus super centrum nucis, pro latioribus ad vel sub centrum nucis. A Gaudichaudia synoptera dif- fert pilis caulis minoribus adpressisque, non paten- tibus erectisve, brachiis trabeculae fere aequalibus in longitudine et numquam subaristatis ut frequent- er in Gaudichaudia synoptera. Gaudichaudia sym- plecta similis indumento caulis, sed pilis caulis mi- noribus saepeque pro ratione latioribus [quam eis G. symplecta|. Formae Oaxacae centralis alis sa- marae reductis a Gaudichaudia cynanchoides se- cedunt pilis parvis v-formibus omnino nullis. Distinguished from lineages in nothosection Cy- clotomopterys, with which it shares a general sim- ilarity in shape of the samara wing, by the narrower postical lobe of the samara wing, by the broader, less acute-acuminate apex of the postical lobe, by the slight to pronounced asymmetry in the lateral lobes of the samara wing, and in having the sa- maras typically widest above the center of the nut, rather than widest at or below the center of the nut. Differing from G. synoptera in having the stem hairs smaller and appressed rather than spreading or erect, with the limbs of the trabecula nearly equal in length and never subaristate as frequently found in G. synoptera. Somewhat similar in stem vesture to G. symplecta, but with smaller stem hairs that are frequently wider relative to their length than those in G. symplecta. In phases from central Oaxaca with reduced samara wings, separable from G. cynanchoides by the complete absence of small v-shaped hairs. JESSUP: GAUDICHAUDIA 251 Paratypes. Mexico. Oaxaca: 15 km W of Oaxaca on hwy 190, 1710 m, 11 Oct 1983, Anderson 13031 (MICH); Mpio. Oaxaca, vicinity of Monte Alban, 21 October 1988, Jessup 4043 (CHAP, IEB, MICH, UC); Mpio. Oaxaca, vicinity of Monte Alban, 21 October 1988, Jessup 4045 (CHAP, IEB, MICH, UC); Mpio. Oaxaca, vicinity of Monte Alban, 21 October 1988, Jessup 4046 (CHAP, IEB, MICH, UC); (CHAP, IEB, MICH, UC). Puebla: 8 km S and W of Tehuacan on rd to Huahuapan de Leon, 1900 m, Jessup 4040 (CHAP, IEB, MICH, UC). Gaudichaudia intermixteca means “‘among the Mixtec,’ reflecting the narrow distribution of this species in central Oaxaca, where it is especially abundant and diverse in the vicinity of Monte Al- ban, but the name also denotes the genetically ‘‘in- termixed”’ nature of this species. The species has clear affinities with lineages in section Tritomop- terys, as shown by phylogenetic placement of the cpDNA haplotype borne by all collections exam- ined. That cpDNA haplotype is, however, unique to G. intermixteca, suggesting an ancient hybridiza- tion that has persisted long enough to evolve a di- vergent genome. Nuclear DNA analysis places the group close to G. galeottiana and the G. albida complex (Jessup 2002). In addition to the concen- tration in the vicinity of Monte Alban, the species is found somewhat further north in Oaxaca and in extreme southern Puebla (Fig. 7). Anderson (1993) found n = 80 pairs of meiotic chromosomes in the paratype Anderson 13031. Nothosection Cyclotomopterys S. L. Jessup notho- sect. nov. (Gaudichaudia sect. Cyclopterys X Gaudichaudia sect. Tritomopterys), Fig. 8. This nothosection includes two new species with re- stricted geographic ranges in the central and western Eje Volcanico Transversal. Gaudichaudia synoptera S. L. Jessup, sp. nov. (Fig. 8)—TYPE: Mexico. Edo. Mexico, Mpio. Tepetilixpa, 3.2 km S of Tepetilixpa on rd be- tween Cuautla and Amecameca, 2160 m, 8 Oct 1988, Jessup 4020 (Holotype MICH; isotypes CHAP. IEB} UC). A Gaudichaudia symplecta et Gaudichaudia in- termixteca secedit pilis caulis longis angustisque, erecto-patentibus ad subaristatis, numquam adpres- sis, caulibus hispidis, brachiis trabeculae inter se valde differentibus in longitudine. A G. andersonii differt samaris ovatis ad orbiculatis sine constric- tione postica alae lateralis prope basin nucis. Separable from G. symplecta and G. intermixteca by the relatively long and narrow stem hairs char- acterized by a marked difference in length of the trabecula limbs, the stem hairs erect-spreading to subaristate, never appressed, and the stems hispid. Distinct from G. andersonii in the ovate to orbic- ular samaras lacking a postical constriction in the lateral wing near the base of the nut. D5, Fic. 8. MADRONO [Vol. 49 Nothosection Cyclotomopterys 2 3 G.synoptera . MS’ D Nothosection Cyclotomopterys fruit morphology and geographic distribution of the holotypes and paratypes. A-E, G. synoptera: A) Jessup 4022, B) Jessup 4023, C) Soto 4024, D) Jessup 4020 (type), E) Jessup 4015; F—H, G. symplecta: F) Jessup 4074 (type), G) Anderson 13291, H) Jessup 4025; 1, G. andersonii: Jessup 4026 (type). Samara illustrations are approximately 1X. Paratypes. Mexico. Guerrero: Soto 4024 (MICH). Estado Mexico: 7 km W of Temascaltepec on rd to Real de Arriba, 2000 m, 10 Oct 1988, Jessup 4022 (CHAP, IEB, MICH, UC); 12 km SW of Temascaltepec on rd to Tejupilco, 1750 m, 11 Oct 1988, Jessup 4023 (CHAP, IEB, MICH, UC). Morelos: ca. | km SE of Laureles village limit, Bar- ranca Tezahuate, ca 3 km NW of Tlayacapan, 1800 m, 6 Oct 1988, Jessup 4015 (CHAP, IEB, MICH, UC). G. cycloptera = Gaudichaudia synoptera is one of two recogniz- able hybrid species formed between species in sec- tion Jritomopterys and section Cyclopterys. It is immediately separable from the other species in this nothosection by the distinct morphology of stem hairs (Fig. 9). The most probable origin for G. syn- optera is as an amphiploid between Gaudichaudia sp. ined. “hirsuta” and G. cycloptera. The evi- dence from morphology and DNA analyses sup- ports that conclusion. The stem vesture is similar 1.0mm C i D Comparison of stem hairs on representative collections from Michoacan. A) G. cycloptera, Jessup 4033; B) G. synoptera, Jessup 4020 (type); C) G. andersonii, Jessup 4026 (type); D) G. symplecta, Jessup 4074 (type); E) G. diandra, Jessup 4034. Fic. 9. 2002] to that found in Gaudichaudia sp. ined. “‘hirsuta,”’ and some of the collections tested share a cpDNA haplotype with members of that lineage. The type bears a cpDNA haplotype that, outside of the G. synoptera collections, is restricted to Oaxaca and Chiapas where it is most typically carried by Gau- dichaudia sp. ined. “hirsuta” and Gaudichaudia sp. ined. “‘velutina”’ (Fig. 1). Nuclear DNA analyses place the type closest to Gaudichaudia sp. ined. “‘hirsuta.”> Other members of the species carry a cpDNA haplotype that is most commonly carried by G. cycloptera. The name means “‘twining together,” reflecting the ten- dency of several twining branches, even from dif- ferent plants, and occasionally from different spe- cies, to form “‘limbs”’ by twining together. The ob- vious analogy is that the lineages themselves are intertwined in this species that apparently originat- ed through hybridization of plants from different sections of Gaudichaudia. The literal meaning of the name and the metaphorical meaning coincide in this plant to the extent that the habit of forming twined limbs complexed from shoots of different genets facilitates wide outcrossing and a reticulated ancestry. The pollinators are anthophorine bees specialized for collecting oil from the calyx glands present as part of the conservative malpighiacean floral syndrome (Vogel 1974; Anderson 1990), and are adapted for the family-level floral characters. The bees likely do not distinguish among species within a genus, and are thus more apt to cross pol- linate divergent lineages when the flowers are closely juxtaposed, as they often are in the tangled thickets inhabited by Gaudichaudia. Gaudichaudia synoptera has a rather narrow geographic distribu- tion (Fig. 8), ranging from southwestern Edo. Mex- ico, where it is especially common in the vicinity of Temascaltepec, eastward into Morelos, and in Edo. Mexico east of Distrit o Federal. Gaudichaudia symplecta S. L. Jessup, sp. nov. (Fig. 8) TYPE: Mexico. Michoacan, Mpio. Mo- relia, 23.5 km E of Morelia on rd from Cd Hi- dalgo, 2170 m, 1 Nov 1988, Jessup 4074 (Ho- lotype MICH; isotypes CHAP, IEB, UC). A Gaudichaudia cycloptera secedit incisura ap- icali insigni alae lateralis et apice acuto-acuminata marginis posticae loborum lateralium alae. A Gau- dichaudia synoptera et Gaudichaudia andersonii differt indumento caulis, pilis adpressis, numquam erecto-patentibus vel subaristatis, caulibus ita seri- ceis, non hispidis, brachiis trabeculae subaequali- bus in longitudine. A Gaudichaudia intermixteca differt lobis alae lateralis samarae plerumque sym- metricis et samaris latioribus ad vel sub medium. Separable from G. cycloptera by the pronounced apical notch in the lateral wing, and the acute-acu- minate apex of the postical margin of the lateral wing lobes. Distinct from G. synoptera and G. an- dersonii in the stem vesture, comprising hairs that JESSUP: GAUDICHAUDIA 29 1oe) are subequal in length of the trabecula limbs, uni- formly appressed, never erect-spreading or subar- istate, the stems thus sericeous, not hispid. Differ- ing from G. intermixteca in the generally symmet- ric lobes of the lateral samara wing, and in having the samaras widest at or below the middle. Paratypes. Mexico. Estado Mexico: 0.5 km N of Amatepec on rd to Tejupilco, 1750 m, 11 Oct 1988, Jessup 4025 (CHAP, IEB, MICH, UC). Michoacan: Mpio. Zitacuaro, Puerto del Gato, 5 km N of Zi- tacuaro on hwy 15, 1800 m, 26 Oct 1983, Anderson 13291 (MICH). The name means twisted and plaited together, re- ferring to the habit, as in the previous species, of forming branches that appear almost braided from separate twining branches. Both the type and one of the paratypes (Anderson 1329]) bear a cpDNA haplotype that is characteristic of section Tritomop- terys. The other paratype (Jessup 4025) bears a cpDNA haplotype that is otherwise confined to lin- eages within section Cyclopterys. Unlike its close relative, G. synoptera, this species has stem tri- chomes that are in all respects similar to those typ- ically found in G. albida sen. str. Nuclear DNA analyses also place members of this species close to lineages within section Cyclopterys. The conclu- sion that G. symplecta originated as an amphiploid cross between G. albida and G. cycloptera is thus well supported. Among the plethora of specific and infraspecific taxa treated by Niedenzu, none is a feasible can- didate for assignment of this species. Niedenzu moved G. arnottiana Juss. to subspecific rank un- der G. pentandra Juss. (=G. cycloptera (DC.) W. R. Anderson) and described three new varieties and two new forms of G. pentandra subsp. arnottiana (Juss.) Niedenzu. One or more of those infraspecific names might be construed as referring to this spe- cies, but the characters Niedenzu emphasized in his diagnoses make it difficult to reach a conclusion about circumscription of his taxa. Jussieu (1843) clearly expressed doubt that his G. arnottiana is distinct from G. cycloptera: ‘“‘admodum. affinis precedenti, cum qua staminibus quinque antheri- feris equalibus couvenit foliorumque forma (fide iconis); calyce, ut videtur, 10-glanduloso ut quibus- dam levioris momenti notis subdissimilis; forsan conjungenda. Species Candolleana forsan conspe- cifica, certe congener fide iconis Flor. Mexic. ined. in qua carpellum calyce eglanduloso stipatum pen- dere e filo videtur.’’ The “‘closely related preceding species” is G. cycloptera (DC.) W. Anderson, and nothing in the protologue clearly differentiates G. arnottiana from G. cycloptera or indicates charac- ters that would suggest the name should apply to the species that I am naming G. symplecta. Jussieu in fact expressed doubt that G. arnottiana is distinct and suggested that they are conspecific. Niedenzu’s infraspecific elaborations notwithstanding, G. ar- 254 nottiana Juss. can be considered a synonym of G. cycloptera (DC.) W. R. Anderson. Gaudichaudia (sect. Cyclopterys) andersonii S. L. Jessup, sp. nov. (Fig. 8)—TYPE: Mexico. Estado Mexico: 1 km S of Temascaltepec on rd to Te- jupilco, 1790 m, 13 Oct 1988, Jessup 4026 (Ho- lotype MICH). A speciebus alis differt samaris magnis incisura apicali insigni alae lateralis, lobos duos laterales symmetricos sinu lata v-forma separatos formanti- bus, marginibus alae lateralis rotundatis et ad con- strictionem sub centrum contractis, lobo postico ex- panso e constrictione sub nucem et interdum cauda brevi abrupta ornato. Distinct from other species in the large samaras with pronounced apical notch in the lateral wing, forming two symmetric lateral lobes separated by a broad v-shaped sinus, the lateral wing margins rounded to a constriction in the wing below center, the postical lobe flared from the constriction below the nut and sometimes abruptly appendaged by a short tail. Paratype. Mexico. Estado Mexico: 1.6 km S of Temascaltepec on Temascaltepec-Tejupilco-Amate- pec rd, 1600 m, 14 Oct 1966, Anderson & Las- kowski 3988 (MICH). This very distinctive species is named in honor of William R. Anderson who has devoted years of field and laboratory work to the study of Gaudi- chaudia, among other malpighs. I found the species during a prolonged foray in the tangled vegetation south of Temascaltepec, occasioned by mechanical failure of the VW microbus, belonging to S. D. Koch, in which Anderson and I were passengers during an expedition into northern Guerrero. An- derson had collected the species in 1966 in the same general area (Fig. 8). Nothing like it has been found anywhere else, and other than the type and paratype cited here, I am unaware of other collec- tions of this species. In vegetative characters and floral morphology G. andersonii resembles both G. cycloptera and G. synoptera. The long, rather narrow stem hairs with distinctly unequal limbs of the trabecula (Fig. 9) are a fairly close match to those found in G. cy- cloptera, but are somewhat smaller than those found in G. synoptera, and although the hairs are distinctly erect-spreading in G. andersonii, they do not form the distinctive subaristate stem vesture found in G. synoptera. The type of G. andersonii was found to carry a cpDNA haplotype otherwise known only from a topotype collection of G. chas- ei. In the nuclear DNA analysis G. andersonii is closely placed with G. cycloptera, well apart from G. chasei, which is placed closer to G. intermixteca and G. galeottiana in that analysis (Jessup 1994). The molecular data suggests that G. andersonii is a hybrid within section Cyclopterys between G. cy- cloptera and G. chasei, and the intermediacy of sa- MADRONO [Vol. 49 mara wing morphology is consistent with that con- clusion. DISCUSSION Tribe Gaudichaudieae, as most recently defined (Anderson 1985) includes Gaudichaudia, Aspicar- pa, Janusia, Camarea, and Peregrina. According to Davis et al. (2001), the tribe is well supported as a monophyletic group nested within a well sup- ported monophyletic stigmaphylloid clade, which includes (in addition to Tribe Gaudichaudieae) Stig- maphyllon and Banisteriopsis, among other genera. A lineage resembling one of the stigmaphylloids is the most plausible ancestor for Tribe Gaudichau- dieae. Examining the distribution of chromosome numbers (Anderson 1993) across the entire stig- maphylloid clade we see in species representing basal lineages a high frequency of nm = 10 meiotic chromosome pairs, probably the base number for the clade. In Banisteriopsis 13 of 14 species re- ported have n = 10 pairs, and one has n = 20 pairs. In Stigmaphyllon 4 of 4 species reported have n = 10 pairs. Within Tribe Gaudichaudieae, ploidal lev- els in Janusia are n = 10 (2 of 12 species reported), n = 20 or, in one case, aneuploid n = 19 (9 of 12 species reported), and n = 40 (1 of 12 species re- ported). Chromosome numbers in Aspicarpa are n = 20 (1 of 6 species reported) and n = 40 (5 of 6 species reported). Camarea all have n = 17, and Peregrina has n = 19 meiotic pairs. In Gaudichau- dia chromosome number is n = 40 or n = 80 (and n = 120 in one isolated collection). The chromosome numbers in combination with the molecular data strongly support the hypothesis that evolution in the stigmaphylloids has proceeded by a series of genomic doublings trending from n = 10 to n = 80, producing a polyploid series on the base number x = 10. In the stigmaphylloid clade, Banisteriopsis and Stigmaphyllon are ances- tral diploids, with a single tetraploid in Banister- iopsis. Janusia is primarily tetraploid with two dip- loid lineages remaining and a single octoploid lin- eage. Camarea and Peregrina are most likely an- ueploid reductions from the tetraploid state. Aspicarpa 1s now understood to be at least in part nested within Gaudichaudia and retains a single known tetraploid lineage, but is primarily octaploid. Gaudichaudia sensu stricto is fundamentally octo- ploid, but developed a series of wide crosses at the sextodecaploid level, including the species de- scribed in this paper. The genus Gaudichaudia is apparently built on an ancient polyploid complex. Regular meiotic pairing among homologous chromosomes resulted in essentially instantaneous diploidization of am- phiploid crosses. Since the trend toward higher ploidy is clearly demonstrated within the stigma- phylloid clade, the obvious hypothesis is that line- ages with lower ploidy will be phylogenetically more basal within Tribe Gaudichaudieae. If that hy- 2002] pothesis is supported by molecular phylogenetic tests, I predict that relict lineages will be found to exist within Gaudichaudia, perhaps as narrow en- demics or isolated populations in central Mexico, that retain the primitive lower ploidal levels. The occurrence of such lineages is already a strong pos- sibility within section Oligopterys, since a single species of Aspicarpa (A. schinnii W. R. Anderson) is known to retain the n = 20 condition (Anderson 1993). That species may or may not properly be- long within Gaudichaudia, and it remains to be seen whether molecular data will place that lineage as basal within the section. A thorough screening of isolated lineages within other sections of Gau- dichaudia is likely to reveal additional instances if indeed any are extant. ACKNOWLEDGMENTS The author thanks William R. Anderson, who gener- ously shared his knowledge and research collections and provided guidance and assistance with field work, and E. Pichersky and R. Fogel, who provided laboratory space and shared equipment. Stephen Koch at CHAPA provided hospitality and generous assistance with fieldwork. Many thanks to reviewers who provided helpful comments on the manuscript. This research was funded in part by NSF grant BSR-8700340 to W. R. Anderson, and by NSF Doc- toral Dissertation Improvement grant BSR-8823076 to W. R. Anderson for S. L. Jessup. LITERATURE CITED ANDERSON, W. R. 1979. Floral conservatism in neotropical Malpighiaceae. Biotropica 11:219—223. . 1980. Cryptic self-fertilization in the Malpighi- aceae. Science 207:892-893. . 1985. Peregrina, a new genus of Malpighiaceae from Brazil and Paraguay. Systematic Botany 10: 303-307. . 1987. Notes on neotropical Malpighiaceae—II. Contributions from the University of Michigan Her- barium 16:55—108. . 1993. Chromosome numbers of neotropical Mal- pighiaceae. Contributions from the University of Michigan Herbarium 19:341—354. BUCHMANN, S. L. 1987. The ecology of oil flowers and JESSUP: GAUDICHAUDIA 259 their bees. Annual Review of Ecology and System- atics 18:343-—369. CAMERON, K. M., M. W. CHASE, W. R. ANDERSON, AND H. G. HILuis. 2001. Molecular systematics of Malpighi- aceae: evidence from plastid rbcL and matK sequenc- es. American Journal of Botany 88:1847—1862. CANDOLLE, A. P. DE. 1824. Malpighiaceae. Prodromus Sys- tematis Natualis Regni Vegetabilis [:577—592. Davis, C. C., W. R. ANDERSON, AND M. J. DONOGHUE. 2001. Phylogeny of Malpighiaceae: evidence from chloroplast ndhF and trnL-F nucleotide sequences. American Journal of Botany 88:1830—1846. Dow LInG, T. E., C. Moritz, J. D. PALMER, AND L. H. RIE- SEBERG. 1996. Nucleic acids III: analysis of fragments and restriction sites. Pp. 249-320 in D. M. Hillis, C. Moritz, and B. K. Mable (eds.), Molecular system- atics, 2nd ed. Sinauer, Sunderland, MA. Farris, J. S. 1989. Hennig86. Port Jefferson Station, New York, NY. GREUTER, W., J. MCNEILL, EF R. BARRIE, H.-M. BURDET, V. DEMOuLIN, D. S. FILGUEIRAS, D. H. NICHOLSON, P. C. Si_vA, J. E. Skoc, T. TREHANE, N. J. TULAND, AND D. L. HAwKSworTH (eds.). 2000. International Code of Botanical Nomenclature (St. Louis Code). Regnum Vegetabile 131. Koeltz Scientific Books, K6nigstein, Germany. Jessup, S. L. 1994. Reticulate evolution in Gaudichaudia (Malpighiaceae). Ph.D. dissertation. University of Michigan, Ann Arbor, MI. . 2002. Reticulate ancestry in Mexican Gaudi- chaudia (Malpighiaceae) analyzed with RAPDs and southern hybridization. Madrono 49:256—273. Jussieu, ADR. 1843. Monographie de la famille des Mal- pighiacées. Archives du Museum d’ Histoire Naturel- le. Paris 3:5—151, 255-616, pl. 1—23. McVauGu, R. 1980. Botanical results of the Sessé and Mocino expedition (1787—1803) II. the Icones Florae Mexicanae. Contributions from the University of Michigan Herbarium 14:99—140. NIEDENZU, F. 1928. Malpighiaceae. Jn A. Engler (ed.), Das Pflanzenreich IV, 141:1—870. SWOFFORD, D. L. 1990. PAUP: Phylogenetic analysis us- ing parsimony, version 3.0. Illinois Natural History Survey, Champaign, IL. TAYLOR, D. W. AND W. L. CREPET. 1987. Fossil floral ev- idence of Malpighiaceae and an early plant-pollinator relationship. American Journal of Botany 74:274- 286. VoGEL, S. 1990. History of the Malpighiaceae in the light of pollination ecolgy. Memoirs of the New York Bo- tanical Garden 55:130—142. MApRONO, Vol. 49, No. 4, pp. 256-273, 2002 RETICULATE ANCESTRY IN MEXICAN GAUDICHAUDIA (MALPIGHIACEAE) ANALYZED WITH RAPD’s AND SOUTHERN HYBRIDIZATION STEVEN L. JESSUP Department of Biology, Southern Oregon University, Ashland, OR 97520 Jessup @sou.edu ABSTRACT Evidence of relationships based on randomly amplified polymorphic DNA (RAPD) data combined with information about cpDNA haplotypes can be used to resolve details of reticulate ancestry in an otherwise intractable polyploid complex in Gaudichaudia (Malpighiaceae). Robust inference of genetic relationships among taxa, however, depends critically on two assumptions: (1) character states compared among taxa are homologous, and (2) characters scored as different are independent. Application of randomly amplified DNA methods, such as RAPD’s, have largely made these assumptions without testing them. In this study I use RAPD’s to elucidate relationships among lineages and to infer reticulate ancestry of amphiploid lineages in Gaudichaudia. | test the assumption that comigrating RAPD fragments are homologous using hybridization of radio-labeled RAPD fragments probed against blots of randomly amplified DNA as an indicator of homology. The probes bind strongly only to fragments on the blots having sequence homol- ogy. Results demonstrate that all gel fragments included in the analysis meet the assumption of homology. Gel fragments can therefore be reliably scored directly as characters. The assumption of independence of RAPD fragments is also explored. Although multiple fragments with strong sequence homology appear in most blots, gel-visualized fragments are generally independent. RESUMEN La evidencia de relaciones basadas en datos de DNA polimorfico aleatoriamente amplificado (RAPD) combinados con informacion sobre haplotipos cpDNA se puede utilizar para resolver detalles, de otra manera insuperables, de ascendencia reticulada en un complejo poliploide en Gaudichaudia (Malpighi- aceae). La solida inferencia de relaciones genéticas entre especies, sin embargo, depende criticamente de dos supuestos: (1) los estados del caracter comparados entre grupos taxonomicos son homdlogos, y (2) los caracteres anotados como diferentes son independientes. El uso de los métodos de DNA amplificado aleatoriamente, tales como los RAPD’s, ha hecho estas asunciones en gran parte sin probarlas. En este estudio utilizo RAPD’s para aclarar relaciones entre linajes y para deducir la ascendencia del reticulado de linajes de anfiploides en Gaudichaudia. Pongo a prueba la asuncion de que los fragmentos comigrantes de RAPD son homodlogos usando la hibridaci6n de fragmentos RAPD marcados radiactivamente como testigos contra manchas de DNA aleatoriamente amplificado como un indicador de la homologia. Los testigos se adhieren fuertemente solamente a aquellos fragmentos dentro de las manchas que tienen homologia de secuencia. Los resultados demuestran que todos los fragmentos del gel incluidos en el andlisis cumplen con la asuncion de la homologia. Los fragmentos de gel se pueden por lo tanto contar confiablemente directamente como caracteres. También se explora la asunci6n de independencia de los fragmentos de RAPD. Aunque los fragmentos multiples con fuerte homologia de secuencia aparecen en la mayoria de las manchas, los fragmentos gel-visualizados son generalmente independientes. Key Words: Polyploid, introgression, Malpighiaceae, RAPD, Southern hybridization. Gaudichaudia (Malpighiaceae) has been revised chromosome numbers, showed combinations of based in part on inference from patterns of variation in cpDNA restriction sites (Jessup 2002). Evidence of relationships among lineages within Gaudichau- dia from morphology, chromosome counts, and amplification of RAPD fragments corroborates the cpDNA evidence and helps to further resolve retic- ulate ancestry of some species in the genus (Jessup, 1994). In particular, a minimum spanning tree anal- ysis of RAPD fragments (based on presence of am- plified fragments visible on agarose gels), con- structed using Jaccard similarity, shows broad con- gruence of cpDNA haplotypes and nuclear ge- nomes. Plants that were inferred to be intersectional amphiploids, based on morphological features and cpDNA haplotypes and nuclear genomes character- ized by RAPD profiles that supported the hybridity hypotheses. While the RAPD data helps define lineages and clearly demonstrates patterns of relationship and ancestry among the lineages, a test of the validity of the underlying assumptions about the RAPD data nevertheless remains worthwhile (Arnold and Emms 1998; Rieseberg 1996; Wolfe and Liston 1998), especially since gel fragments alone might be useful for inferring relationships. Homology of character states and independence of characters are prerequisite features of characters used in many systematic analyses. Restriction fragments are 2002] strongly correlated, which is why restriction frag- ments alone cannot be scored for presence/absence and used directly as characters. While randomly amplified DNA markers, such as RAPD’s, have not been shown to be correlated in the same way that restriction fragments are, estimates of relationship based on RAPD fragments would be similarly skewed if tightly linked markers were treated as independent estimators of relatedness (Lynch 1988). Estimates of relatedness would also be in- accurate if fragments of the same electrophoretic mobility were commonly of heterologous origins. In this paper I present the results of the RAPD anal- ysis in Gaudichaudia and report experiments that test RAPD band homology and independence using hybridization of *?P labeled RAPD probes to south- ern blots of RAPD gels. I demonstrate the utility of the procedure in identifying the specific ancestry of amphiploids. When combined with knowledge of cpDNA haplotypes in the samples, this proce- dure further resolves reticulate ancestry by speci- fying which of the parental lineages contributed the maternal genome. Expected behavior of RAPD markers in amphi- ploids. In tetraploid Gaudichaudia where chromo- somes from divergent lineages may reside within the same nucleus, and where the lineages are re- producing primarily by selfing (Anderson, 1980), markers that occur within the same set of chro- mosomes can be tightly linked. When homeologous chromosome sets are divergent but still pair at mei- osis in wide amphiploid crosses, we expect a marked reduction in viable gametes and reduced fecundity in the F, progeny. Given that strongly in- breeding lineages quickly approach fixation of var- iable alleles (Li 1976), it seems likely that a pre- ponderance of markers will frequently be fixed within a set of chromosomes in strongly selfing lin- eages. With little or no recombination between homeologous chromosomes most variability occurs among rather than within homeologous sets of chromosomes. Lineages with size variants of the same marker fixed on different (homeologous) sets of chromosomes—synologous loci as defined by Mindell & Meyer (2001)—will exhibit fixed het- erozygosity. The variants that occurred in the an- cestral diploids as orthologous loci, as fixed differ- ences at the same locus in different lineages, are combined in the amphiploid as synologous loci, though they are less likely to be fixed in progeny resulting from outcrossing. Since RAPD sites are restricted to individual chromosomes, they neces- sarily obey all of the constraints associated with chromosomal inheritance. We expect that RAPD fragments of the same mo- lecular weight—fragments that appear at precisely the same location on a gel—are products of ho- mologous sites. There is, however, the possibility of heterologous fragments with the same mobili- ty—tfragments that are identical in state but not ho- JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 257 mologous. We also expect that fragments of differ- ent mobility are independent products of unlinked sites. The assumptions of homology and indepen- dence, then, can be reduced to two questions. (1) Is there sequence homology among bands of dif- ferent electrophoretic mobility within a sample? If we count multiple bands as distinct when they are in fact the result of a single locus then we violate the assumption of independence. (2) Is there se- quence homology among bands of the same elec- trophoretic mobility among samples? If we count bands of the same electrophoretic mobility as iden- tical when they are in fact distinct then we violate the assumption of homology. Testing those assump- tions is simple in principle. An estimate of se- quence homology can be obtained by observing the relative strength of hybridization signal between a labeled RAPD probe of known origin and samples from bands of the same molecular weight whose homology is in question. Bands that hybridize strongly to the probe are inferred to have a high level of sequence homology with the probe. By in- ference, a strong hybridization signal is an indicator of close relatedness. MATERIALS AND METHODS DNA extraction. Plants in the genus Gaudichau- dia were sampled from a wide geographic area in Mexico, representing the center of diversity in the genus. Detailed source data for the collections is reported elsewhere (Jessup, 1994). Total genomic DNA was extracted from leaves that were snap- frozen in liquid nitrogen and subsequently stored at —80°C. Total genomic DNA extraction was based on the CTAB procedure presented by Hillis et al. (1990) and Dowling et al. (1996). All DNA extrac- tions were purified by ultracentrifugation on cesium chloride gradients, dialysis to remove the cesium chloride, and ethanol precipitation following pro- cedure in Sambrook et al. (1989). Final yield of DNA was between 25 and 750 pg DNA per ex- traction (from 1—2 grams of leaf tissue). In all, 134 different DNAs were successfully prepared for use in the molecular procedures. DNA concentrations and estimates of DNA purity were calculated from optical density measurements at \ = 260, A = 280, and A = 320 nm, on a Beckman DU-64 UV spec- trophotometer. RAPD reactions and Southern hybridization. Pu- rified template DNA was diluted to 1 ng/ml in a reaction elixir buffered to pH 8.3 with 10 mM Tris- HCl containing 50 mM KCl, 2 mM MgCl, dNTPs at a concentration of 100 M each, five arbitrary 10-base oligonucleotide primers (Operon®) were used in separate reactions, each at a concentration of 0.2 ~M, and Taq DNA polymerase (Perkin E]- mer®) was used in all reactions at a concentration of 0.07 units/ng template DNA. The reaction was carried out in a total volume of 25 wl on an M.J.Research® thermal cycler programmed at max- 258 imum ramp speed for: 1 cycle of 30 seconds at 94°C; 45 cycles of 1 minute at 94°C, 1 minute at 35°C, and 2 minutes at 72°C; and 1 cycle of 5 min- utes at 72°C, followed by a hold at 4°C. The products of the PCR reactions were separat- ed by electrophoresis on 2% agarose gels with 200 ng/ml ethidium bromide, at 25V constant voltage for approximately 14 hours at room temperature in 1X TAE, pH 7.6. A 100-bp ladder (Pharmacia Bio- tech®) was loaded into three or four lanes on each gel at even intervals among samples as a high res- olution molecular weight marker. Each gel run con- tained from 120 to 134 sample lanes and 12—16 molecular weight marker lanes over four gels. RAPD fragments were visualized by UV transillu- mination, then photographed on Polaroid-57 at f8, 3’ 20”. Negatives were washed in NaSO, 18% for 1 minute, then rinsed 30 minutes in H,O. RAPD fragments visible on the negatives were scored by careful measurement on a lightbox, and molecular weights were determined with the gel analysis pro- gram, Gel Match®® (UVP). All visually detectable fragments on either the films or the scanned images were scored as present. Computer image enhance- ment was used to intensify faint bands. Lanes not showing a fragment at the same position on the gel were scored as absent. RAPD fragments from the PCR reactions were collected from ethidium bromide stained gels under UV transillumination. Bands selected for **P label- ing were sampled by inserting the tip of a pasteur pipet into the center of the band and applying light vacuum pressure with a pipet pump as the tip was withdrawn from the gel. This produced a cylindri- cal gel section about 1 mm wide by about 7 mm long while preserving the source gel for blotting. The gel section was extruded into a microfuge tube and kept on ice. Collected fragments were diluted 10:1, reamplified using the original PCR protocol, then electrophoresed on 3% low Ty, agarose gels to further purify the fragment. The whole reamplified fragment was collected after gel purification and 12 pl was radio-labeled with a-**P tagged ATP using a polymerase reaction. A few ng of molecular weight marker were also labeled at the same time as the RAPD probes. La- beling reactions were carried out using random priming with a mixture of hexadeoxyribonucleo- tides (6 bp oligonucleotides) according to the pro- tocol developed by Feinberg and Vogelstein (1983, 1984). This technique resulted in probes labeled to high specific activity. Unincorporated nucleotides were separated from labeled RAPD fragments us- ing sephadex columns set up in 9” glass pasteur capillary pipettes. Purified labeled probe was de- natured by immersion in boiling water for 10 min- utes, then ‘quenched’ on ice for 3—5 minutes before beginning the membrane hybridization reaction. RAPD gels were blotted to nylon (Zetabind®) membranes following procedures in Maniatis et al. (1989). Membranes were allowed to dry after trans- MADRONO [Vol. 49 fer of the amplified fragments then stored at room temperature until Southern hybridization. Prehybridization and Southern hybridization re- actions followed procedure outlined in Dowling et al. (1996). The prehybridization solution was 4X SSC, 1% SDS, and 0.5% nonfat dry milk. Blots which had not been previously probed were first pretreated by washing in 0.1X SSC, 0.5% SDS for 1 hour at 65°C. Prehybridized blots were removed from the incubator and all but about 15 ml of the prehybridization solution was removed from the hybridization tray. The labeled probe was then add- ed to the tray and thoroughly mixed. Several blots were hybridized simultaneously. Care was taken not to introduce bubbles into the space between blots. Hybridization reactions were allowed to pro- ceed for 12—18 hours at 62°C. Hybridized blots were removed from the trays and washed in three or four changes of 2* SSC, 0.5% SDS: two short washes at room temperature followed by one or two 30—45 minute washes at 60°C. Blots and discarded wash were monitored with a Geiger counter during the wash procedure to assess when background radiation on the filters had been adequately reduced. Filters were removed from the final wash and blotted to remove excess wash, then wrapped in plastic and placed in x-ray film cassettes with intensifying screens. Kodak X- OMAT® AR film was loaded into the cassettes and they were exposed over night at —85°C. Films were removed and developed on an X-OMAT® auto- matic X-ray film developer. Following autoradiog- raphy, blots were stripped of probe in hot 0.5 SSC and monitored until radiation was reduced to low levels. Blots were then prepared for reprobing with a different RAPD fragment. RESULTS Relationships inferred from RAPD fragments. Seventy-five plants were scored for 79 RAPD sites. Table 1 records the molecular weights of RAPD fragments scored from the gels for each of the primers used in the study. Table 2 records the frag- ments scored from the gels for each collection. Us- able sites were those that could be consistently scored for all 75 collections included in the analy- sis. Figure 1 shows a typical RAPD gel, in this case with RAPD bands produced using primer sequence CAAACGTCGG (A-19). Collections are arrayed on the gels by taxon defined on morphological sim- ilarity, and within taxon by geographic region. There are four marker lanes per gel, each a 100 bp ladder with molecular weight indicated for the 800 bp fragments. Blank lanes are collections that did not amplify. Blank lanes are excluded from the data tables. Notice the general pattern of shared bands. Bands of the same molecular weight tend to occur in adjacent lanes on the gel, i.e., among collections grouped by taxon and geographic proximity. The data were clustered with a minimum spanning 2002] TABLE 1. RAPD FRAGMENTS AMPLIFIED FROM 76 COLLEC- TIONS. Size of each fragment (kb) is reported for six dif- ferent primers used in the analysis. Column numbers refer to the data matrix (Table 2). Column 1-16 17-31 32-34 45-49 50-62 63-79 Primer A-9 A-10 A-15 A-16 A-18 A-19 kos ~ 228) 2.60 3.50. — 1-95, 3.70 ys 2et5 S90) 32:00: — 1.70-—'3.50 Eos -- 2:00 12:80.> 1-90 — “1.607 2.50 1.55 1.65 1:60" 1:40~— 1.53 °°'2.20 1.45 60> 50) -" 30" ‘1.45 1.90 £35 P50 3t55 L420 © 1270 PaG, — E30\<- £25 138 1.50 5 £20... £20 L395 1.48 E20 2 Ris. ero 1.28 1.45 Lal fe | 1.00 1.20 1.43 ROS ~- 0:95-0:90 1.15 1.40 95; 0:85-0:80 1.00 1.38 ESS | O70 S “O75 O90 54230 0.80 0.65 1.20 0.70 0.60 1.10 0.60 0.90 0.85 tree using the program Minspan (Podani 1993). A minimum spanning tree is the branching graph of OTU-wise association coefficients that minimizes the sum of all edges. Minimum spanning trees were computed for the RAPD data using the coefficient of Jaccard (Sneath and Sokal 1973), which does not include negative matches as a component of simi- larity (or dissimilarity). This is necessary when es- timating relationships from randomly amplified DNA data since absence of a site does not convey any useful information about relatedness. Many dis- tantly related OTU’s will have state 0 for a large number of sites. Most of those characters, identical in state, will not be identical by descent. The RAPD minimum spanning tree is plotted (Figs. 2—4) and mapped with representative fruits from plants bear- ing each of the cpDNA haplotype defined in Jessup (2002). Each plant included in the analysis is des- ignated with an OTU number in the diagrams cor- responding to a row in Table 1. Correspondence of cpDNA haplotypes and RAPD’s. The longest span in the tree falls between OTU 3 and OTU 5, and effectively defines a left and right half of the minimum spanning tree. cpDNA haplotypes from section Tritomopterys map in part to each side of the minimum spanning tree (Fig. 2). Most plants bearing cpDNA haplotypes from section Tritomopterys with asymmetric fruit wings map to the right half of the tree. All but one plant (OTU 5) bearing cpDNA haplotypes from section Tritomopterys that map to the left side of the tree have irregular or intermediate fruit wing symmetry and belong to G. implexa Jessup (notho- section Tritomochaudia). Plants bearing cpDNA haplotype D (OTU’s 64, 66, and 76) or cpDNA JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 259 haplotype B (OTU 59) having large symmetric fruit wings typical of G. cycloptera (DC.) W. R. Ander- son map in a cluster far to the right with plants from section Tritomopterys and are all members of nothosection Cyclotomopterys, either G. synoptera Jessup (OTU’s 59 and 66) or G. symplecta Jessup (OTU’s 64 and 76). Plants with asymmetric fruit wings bearing cpDNA haplotype D (OTU’s 2, 3, 6, 8, 18) are all members of section Tritomopterys and map just to the right of the span separating the two halves of the spanning tree. Plants with cpDNA haplotypes from section Cy- clopterys (OTU’s 21, 59, 61) map to the same area on the minimum spanning tree as those with similar fruits bearing cpDNA haplotypes from section Tri- tomopterys (Fig. 4). Plants in G. mcvaughii W. R. Anderson, however, map to the far left (OTU’s 22 and 23), even though they carry cpDNA haplotypes that clearly place them in section Cyclopterys (Fig. 4). In particular, OTU 21 (in G. cycloptera), far right, and OTU 23 (in G. mcvaughii), far left, both bear cpDNA haplotype AB from section Cyclop- terys. One plant with morphology of G. cycloptera, and bearing a cpDNA haplotype from section Cy- clopterys (OTU 19), maps closer to the asymmet- ric-winged plants bearing section Tritomopterys cpDNA haplotypes. Plants bearing section Gaudichaudia cpDNA haplotypes all map on the left side of the RAPD minimum spanning tree (Fig. 3). One plant bearing a section Gaudichaudia cpDNA haplotype but with fruit wing morphology similar to G. cycloptera or G. mcvaughii, maps to the far left close to the po- sition of G. mcvaughii. Several plants having inter- mediate fruits and carrying section Gaudichaudia cpDNA haplotypes (Fig. 3) map just to the left of comer (OOS 52. 67, 5608;.09; 70. 71, 75). Plants with similarly intermediate fruit wing shapes but carrying section Tritomopterys cpDNA haplotypes map in the same region on the RAPD tree (OTU’s S49, 50, 56, 655° 72)2 All of ‘the plants with intermediate fruit wing morphology mapping to this region of the minimum spanning tree are G. implexa Jessup in nothosection Tritomochaudia. Plants with morphology typical of G. cynanchoides H. B. K. map near the center of the left half of the tree (OTU’s 40, 41, 42, 44, 45, 46, 47, 48). Three plants with intermediate fruits map to the far left, one bears section Tritomopterys cpDNA haplotype L (OTU 50), and the other two bear section Gau- dichaudia cpDNA haplotype S (OTU’s 49, 51). Those plants are also G. implexa. Plants in G. galeottiana (Nied.) Chodat, or with fruit morphology approaching that of G. galeotti- ana, and bearing section Zygopterys cpDNA hap- lotypes (Fig. 4) cluster together on the RAPD tree on one long branch just to the right of center (OTU’s 27, 28, 29). Plants in that cluster with slightly asymmetric wings are G. zygoptera Jessup (OTU’s 28 and 29) or G. intermixteca Jessup (OTU 26) in nothosection Zygotomopterys. Only one MADRONO [Vol. 49 260 OOOTOOODDOTOOLOOODDOTOTOOOOTLELOOODOTOTLOOOODDNN0TOOTOLTOTOOOODOTOLTLOOONOODOTOOO00TOO0N LV SLOVE uesvoysli OODOOTLOOODTOOTOTOOOOOTOOOTLOLOOOTOLOOOTLLOONONOOTOOTOTOOOTLOOTOTITLTOLOOOONLOO000000 v9 VLOVE ueoeoyoly OOOTLOOTOOOOOTOOLTOOOTOLOOLTOOTOOOTOOOOTLOOOOOTLOOOLOOOTOOOTOTOOTOOTLTOOLTLOOLOOLOO CL Leort UBsBOY STA OODDDD0 TOO TOOODDDDNDNDNNDND000000000000000 LODDD0D000DDDDDOT LODO TO LOOOO0O0D000LLOO000000 VL 9COrl uBdBOYTIAT OODDDOTLOOODODOODOOTOTLOOOOOTLTOOOOOTOOOOTTOOODOTTOOOTTTOTOOOTOTLIOOOOTTLOTOOOOTLILO ell ceOrt uesvoyoly OODDDOTLOOODDDOTILOTOOOOLOODDDDOTOOOOTLOOOOOTLIOOOTTILLIOOOLOLOOOOTOLTOLOLOLOLOLLO [E 60EE1V uesvoyolyy ODDDDDDDD0D0D0000000DDDTOLOOODDTOOTITILOOLLOOOOOOTOTLTOTOLELOOOOOTLOOOOLOLOOLOON000 8 96CL IV uesvoyolyy OODDOLOLOOOTLOOOLOOODDNTOODDOTOLOODDNDNDDDDDDNDNN000000TOODDTOOTOOTOOTOOTOTOLTLOOO00000 OL l6cclV uesvoyol|| OOOLTLOOODOTOOTOOOODODDDDDDDNDNDLOLTOLTILTLOOLOOONNNOTOTLOTOOLOOOOTOTLOOOOLOLOOONDNLOOON OV O9CC8A OSTXOIN OOTOTLOLOOTOOTOLOODDDDDD00TOTOOOTOOODDTTLOOODDDD0TOOTOOTOOOTOTLOOLOOOTLOOOO00N00 ©9 9COrL OOTXOT OOOOTLOLOOTOOTOLOODDDDDDN0TOOODODOTOODOOTTLIOOOONDDDDTOOTOOTOOOTOTTOTLOOOTLOOO00000 c9 ScOvE SOPXSTNI OOTOTLOOOOLOLOODDDDDDDDDD0000000000000TOTOODDNDDDDDD00000000000TODDO0TOO000000000 19 ecOrt COROT OOTOLLOOOOLOLTOODDDDDDDDD00TOODDOTODDDNTOTOOODDDDDDTOOTOOTOOOTOTLOOLTOOOTLOOOO0000 09 1cOvt OSTXOT OOOOTOOOLOTOOTOTOOOODNDDDN0DNDD000NDTOOODDOTTLTOOTOOOOOTOOTLOOTOOOOTLOOTOOOTLOO000000 6S OcOrt OSTXOTN OOODDDOTLOTOOOOTTLOOOODOOTTOTOOOOTOOQOOTOOOOOTLOOOTLILLOLTLOLOOOOOOTLLOLOTLOOOON ie CLELIV OoIxo| OOOLOOLOOLOOLOODDDDDDDDDDDTOODDDOTOODDTOTOOLOOODDDTOODDTOOOODNDDOTOODODOTOOOTOO00000 Ic c80rl Ooster OODOTLOOOOTOOOTOODDDTOLOOOOOTOOOOTLOOOTOOOODDDNDDDOTOOTOTLOTOOTOOOTTOLOOON0000TON0N WL c80V£ OosTTes OODOLTLOOOOLOOQDDDDDDNTOLOOODOTOODDDDTOOTOOOODOOTOOTOOTOTLOTOOTOOODDOTOTOOOOTLOON OL 180rf OosT[ef OOOOTLLLOOTOLOOOTOOOTOLOOOOOTOOOTTELOLILOOOOTLOOTLOOLOOOOTOOTOOOOOTTOTOOTLOLOOO 69 O80rl oosTyef GCOOOLELLOOLOOOLOLOOOLOLOOLOOLOOOLOOLOLLOOOOOLLOOOLOLOOLLOLOLOOLOOOLOOLOOLOLEL LO 89 6LOVL SOSTIE | OOOLTLOOOOTLOOLOOOOOOTOLOOOOTLOLOOLTLTLOOLOOONOTOLTOOTOTOLLOOOOTOOOOODODOTOTOOO00TONN CV SLOVL Costes OOOLLOOODODLOOTOOOOOOTOLOOODOTLILIOOTTLOOLLOOOOOOTOOTOTOTLLOOONOOTLIOOOOTOTOOOOTLOOO vv LLOVE OosTIeL OODOOTLELOOLOOOOOTOOOTOLTOODNDTIOOOOOTOTLLOONNODODOTOTOOTLOLOTOOLTOOOTOOTOOOOTLONN L9 9LOVE oosT[ef OODDTOOTLOTOOOTLOONDDDDDDNDN00000TOTOODTOODDDTTOOOTOTOOTOTLOOOOTOOOTOOODDNDOTLLONN GIL LOLEV oosTyef ODDDCO0D0D0D0D00000000000TOTOOOOOTOTOTLIITLOOTOONODDDOTOOTOTOLOTOOTTLOLTOOLOOO0000000TO000 CV CLIVE os]epiH ODODDDDDD0D000000000000TOLOOODOTLOOOTTTLOOLLOOOOOODODOOTOTOTOTOONDOOTOOOTOOTOOOTOTONN IV ITIv£ osyepry OOOTLOOODDLOOTOOODODOTOLOOODNTLOOTOTTOOLLOONOOOTOOTOTOTOTOONNOOTLOOOOTOTOOOOTLONO OV 601 rf osyeply OOOTOTLOOOLOOOOOTOOOTOTOONDDODOOTOTTOLLOOTOOTOOOOTOTOTLOOLOOOOTOOOOTOTOOOOTIILOT cS ccScead o3|[eplH ODDDD00000000TODDOOOTOLODDDDTODNNN00000 LODDDD000N0TOTOOO00D0000000000000T000000000 6£ LOIVE oyenfeurny OOOLTLOOLLOOOOOOOTOOTLOLOOODDLOOOOTLTOOOLOONDDDDDDOOTOTOLOTOOOTOOTOOQODDNTOOTOONTOOD EC SICC8>I CTS) ODDDD00 LODDD0TODDDDDDDDDN0000000TODDDDTODDD000000TODDDDD000000000TODDDD000TLONN OC OISTV O1OIIOND) OOOTLLOOOTLOLOONNDNNDDDDDDDD0D0DNNDNNN00LOLODDDDDD0000TOOODO00000TLOOODODOTOOTO000 T0000 O€ 690rf ONSEBTMS OODODDDOTLOLOODOTLOOOO0D0D0DDD00000NTOOOOOLODDDDTLOODDND00000000000000TTTOTOTOTOONN G L9OOVE Oe) OODODDDOTLOLTOOOOOTLTOOODDOLOOOODOOTOOOODOTOOOOOTLIOOOTTILELOTLIOLOOOOTOTLTLOLTOLOLOONN 9 990rf OSTOMY) OODDDOTTLOLOOOTODODDDDDDDD00000TOOOTLOODNDNDD0000TOTOOTLTOLTOONOOOTTOOQDDNDD000000TO0 61 6cOrl OTSIIONE) OOOLOLOLOOTOOTOOOOTLLOOLOOODNDNOOOTLOOOLOOODODOOTIOLOOOTLLOOOTOOLOOOOTOTOOTTOOON00 GE 669CI1V BuNTTO) OODOODD0OLTLOTOOOOTOLOOODDODTLOOOOTOLTOOOOOTOTOOOOOTOOTLIOTLOOOTOTTOTLITLTOOOOOTOTOOOOTO /E\| 190rf sedeiy) ODODDDDD0LOLOOOOTOODDDDDDLODDDDOOTOODDDLOLOOOOOOOOTTOTLOOOTOTLOTILOOOOTLOLOONNN0 91 O90rf sedeiy) OODDDD0D0D00TLODDDNNNN0DDDDDDDDDNDNDNDOTOOODDTOLOOOOOOOOTITLLOOOTOTOOOTLOOOOTTOTOOOTOO cl 8SOrl sedeiy) LOOOOOOTLELOONDNNDN0000000000DN0NNN000000 LODDDDDNDN0N00000TODDDDDDNDNDO00TOODOOTOOOTOTOT I 9SOrL sedery) OODOOTLTOONDDTODDDNDOTOTOONDDDDDDDNDNOTOTOOOTOODDDNDNDOOOTLIOLLOOOLOOLOOOTOOODDDD0D0TOOTOT IT SSOrt sedeiy) OLTOOOOTOTLOTOOOOTOLOONDDDDDDDNDNDNTOOODDLOODDDDDOOTOTOTLOTOTOTOLOOOOOTOOOTTLOOTILO el SCCEIV sedery) OLTOOOOOLOTOTLOOOOLOOOOTLOOOOTOOOLOOODDLOODDDDNDDOOTLIOTTOOOTLTLOTOTLOOOOOOOTOOOTLO cl VCCEIN sedeiy) OL 09 OS OV O€ O? 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Ais 800 bp# 7 Su 102 104 106 108 110 112 113 115 117 119 121 123% 24 126 128 130. 132. 134 Fic. 1. ie Example of RAPD products on agarose gel, in this case amplified with primer A-19. Gels were stained with ethidium bromide and visualized with UV transillumination. Molecular weight marker fragments are 100 bp apart. Lane numbers and the 800 and 1400 bp marker fragments are labeled. plant bearing a section Zygopterys cpDNA haplo- type (OTU 57) maps on the left side of the tree. That plant is notable in the morphological similarity of its fruit to other plants with intermediate samaras that carry section Gaudichaudia cpDNA haplotypes and map to the same region of the RAPD tree (OTU’s 52, 68, 75). It also falls close to OTU 58 which has similar fruit shape but bears section Tri- tomopterys cpDNA haplotype L. OTU 57 is a geo- graphically isolated plant carrying a section Zyg- opterys cpDNA haplotype. A plant from the type locality of G. chasei W. R. Anderson (OTU 24), carrying cpDNA G. chasei cpDNA haplotype AE, maps close to the cluster of G. galeottiana on the RAPD tree (Fig. 4). A plant with intermediate fruit wing morphology (OTU 26), bearing section Tri- tomopterys cpDNA haplotype A, subtends G. chas- ei on the RAPD minimum spanning tree (Fig. 2). The only other plant collected that carries G. chasei cpDNA haplotype AE (OTU 63) is G. andersonii Jessup which maps with plants in G. cycloptera carrying section Cyclopterys cpDNA haplotypes (OTU’s 21, 61, 62). The narrowly endemic sub- shrubs, G. krusei W. R. Anderson (OTU 30) and G. 2002] JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 263 RAPD minimum spanning tree mapped with fruits from plants bearing section Tritomopterys cpDNA 23) Qs apy 2060-49) 89) 88) 87474044 — 67) 68 a) @2) 4348) G6) 70)69 ) haplotype OTU(s 42) Se GB. YA A 26 (G. intermixteca) Wy We tly B 4 (G. albida) 15, 16, 17 (G. ‘velutina’) e iy B 59 (G. synoptera) C 11 (G. ‘intermedia’) D 2,3 (G. albida); 6, 8 (G. diandra); 18 (G. ‘hirsuta’) D 66 (G. synoptera) D 64, 76 (G. symplecta) F 1 (G. albida) K 12, 13 (G. ‘intermedia’) L 5 (G. albida); 7, 9,73 (G. diandra); 14 (G. ‘intermedia’) L 25, 50, 53-56, 58, 65 (G. implexa) M 72 (G. implexa) N 10 (G. diandra) 28) B5) Fic. 2. Samaras from plants carrying cpDNA haplotypes from section Tritomopterys plotted on the RAPD minimum spanning tree. subverticillata Rose (OTU 31), map together on the extreme right side of the RAPD tree (Fig. 4). Those taxa are similar in the cpDNA haplotypes they car- ry as well. Gaudichaudia subverticillata and G. krusei form the paraphyletic section Archaeopterys comprising the internal outgroups to section Tvito- mopterys in the cpDNA haplotype phylogeny (Jes- sup 2002). OTU 38, which also carries a cpDNA haplotype positioned as an outgroup to section Tri- tomopterys in the phylogeny, has a fruit shape sim- ilar to G. cynanchoides and maps on the far left of the RAPD tree, close to G. mcvaughii. Southern hybridization results. Table 3 presents the results for lanes on the blots that showed hy- bridization to the probes. The number of fragments per lane hybridizing to the probe is given in the table for each probe used in the study. Chloroplast DNA haplotypes determined with restriction frag- ment analysis (Jessup 1994, 2002) are also indicat- ed for each lane in Table 3. In each of Figs. 5—10, the probe source is represented by an illustration of a samara at the top of the figure or to one side of the autoradiograph. A line points to the fragment that was labeled for the probe. Superimposed on the image of the autoradiograph are illustrations of Samaras from plants that showed strong hybridiza- tion to the probe. The results thus presented show the intersection of data from RAPD’s, cpDNA, and fruit wing morphology. When blots of the RAPD gels were probed with a labeled 1400 bp fragment from lane 12 (Anderson 12937, G. diandra (Nied.) Chodat, Morelos, lane 12) all (100%) of the plants expressing the 1400 bp fragment on the gel showed strong hybridization signal (Fig. 5). The surprising result was that all (100%) lanes showing hybridization to the probe hybridized to fragments of different molecular weights as well. Not surprisingly, the 1400 bp frag- ment (probe source) on the blot showed the stron- gest hybridization signal. Other fragments showing strong hybridization signal (secondary bands) on the blot showed up on the original gel as faint bands, but none of the secondary bands hybridizing the probes were visible on the gels as bright bands, and some of the secondary bands were not at all visible on the gel. Only rarely (one in twenty) did a lane without a 1400 bp fragment show hybridiza- tion to a fragment of a different size (middle frame, bottom panel), and that hybridizing fragment was the same molecular weight as a secondary fragment in lanes with multiple fragments hybridizing to the probe (Fig. 5). The same pattern of results is re- peated with the other probes. All bands of the same electrophoretic mobility hybridized to the probe 264 MADRONO [Vol. 49 RAPD minimum spanning tree mapped with fruits from plants bearing section Gaudichaudia cpDNA 23 Oe ay SNS "Ds v 4) 4S IS a S ? A OE) Ee Se WW. W Fic. 3. 43.49 a8 COGN Ga 40, 41, 44-48 (G. cynanchoides) 49, 51, 67-71 (G. implexa) 42 (G. cynanchoides) 52 (G. implexa) 75 (G. implexa) F 20 (G. cynanchoides x cycloptera) H8OGOSOL Samaras from plants carrying cpDNA haplotypes from section Gaudichaudia, plotted on RAPD minimum spanning tree. The right side of the tree is omitted since plants carrying section Gaudichaudia haplotypes are restricted to the left side of the tree. made from a band at that location. This clearly demonstrates that bands of the same molecular weight have sequence homology and can therefore be reliably considered identical by descent, at least in Gaudichaudia. Oligopterys 32 (G. hirtella) Fic. 4. chaeopterys, and section Oligopterys. RAPD minimum spanning tree mapped with fruits from plants bearing cpDNA haplotypes from sections Cyclopterys, Zygopterys, Archaeopterys, and Oligopterys The probe source in Fig. 6 is G. diandra. All of the plants hybridizing to the 1400 bp probe are found to cluster with the G. albida complex (sec- tion Tritomopterys and intersectional hybrids with section Tritomopterys) in the RAPD minimum AQ o@ CSS, cb AS 66/30-31) O-O—O0-8'@- aye haplotype _ section OTU(s) 73) (9) 42) (1) 41) Y Cyclopterys 19 (G. cycloptera) AB Cyclopterys 60, 21 (G. cycloptera) 49 43 AB Cyclopterys 61, 62 (G. synoptera, G. symplecta) AE Cyclopterys 24, 63 (G. chasei, G. andersonii) AB Cyclopterys 23 (G. mcvaughii) AC Cyclopterys 22 (G. mcvaughii) @7) 26 34 AG Zygopterys 27 (G. galeottiana) 14 ip AD Zygopterys 57 (G. implexa x zygoptera) ata 4) 6 Ww Zygopterys 28, 29 (G. zygoptera) NG & ; bees HC G Archaeopterys 38 (indet. amphiploid) Spl | I Archaeopterys 31 (G. subverticillata) () 28 (is ee) J Archaeopterys 30(G. krusei) Samaras from plants carrying cpDNA haplotypes from section Cyclopterys, section Zygopterys, section Ar- 2002] spanning tree (Figs. 2—4). The lanes showing hy- bridization to the G. diandra probe represent plants carrying cpDNA haplotypes from sections T7vito- mopterys, Cyclopterys, and Zygopterys. All of those plants bearing cpDNA from other than section 77i- tomopterys were identified as amphiploids on the RAPD minimum spanning tree. In the hybridization shown in Fig. 6, the probe (1150 bp) was taken from a member of the G. cy- nanchoides complex (Jessup 4112, Hidalgo, lane 75). The probe was hybridized against the same set of blots shown in Fig. 5 (i.e., from primer A-19 gels, Fig. 1). Again, all of the lanes showing hy- bridization signal except one are clustered together on the RAPD minimum spanning tree (Fig. 3), in- cluding plants identified as G. implexa, hybrids with maternal G. albida and paternal (pollen donor) G. cynanchoides. One lane, representing a plant with a samara morphology diagnostic of the G. al- bida complex (lane 6, Jessup 4052, Oaxaca), shows strong hybridization with the G. cynanchoides probe. That lane represents the same plant found clustering anomalously with the G. cynanchoides group on the RAPD minimum spanning tree (Fig. 3, OTU 5). The only other lane represented by a G. albida samara (lane 119, Jessup 4032, Michoa- can) shows a relatively weak signal, suggesting some involvement with the G. cynanchoides group, perhaps via introgression. The two strong signals in the right panel of the top gel (Fig. 6) are bound probe from the first run (cf. Fig. 6) that did not melt from the blotted DNA during stripping of the probe. These “‘ghost band”’ appear on other blots as well. The hybridization run represented in Fig. 7 used the same blots discussed above. The probe for this hybridization experiment (1900 bp) is from the same source as that described for Fig. 6 (Jessup 4112, Hidalgo, lane 75). Here again we see mainly plants from the G. cynanchoides group hybridizing to the probe (most of gel panel 3). Again we see the anomalous plant from the G. albida complex (lane 6, Jessup 4052, Oaxaca) hybridizing to the probe. A possible G. cycloptera X G. cynanchoides hybrid also shows up on this autoradiograph (gel panel 2). Aspicarpa, the putative sister lineage to Gaudichaudia, has recently been shown to nest, at least in part, within Gaudichaudia (Davis et al. 2001). The Aspicarpa accession used in this study (G. hirtella (Rich.) Jessup, formerly in Aspicarpa), also hybridized to the 1900 bp probe. Three of the suspected amphiploids (G. implexa) hybridizing to the 1900 bp probe (gel panel 4) also hybridized to the 1150 bp probe (Fig. 6, panel 4). Two of the amphiploids (G. implexa) hybridize to the 1900 bp probe but not to the 1150 bp probe, and two (also G. implexa) hybridize to the 1150 bp probe but not to the 1900 bp probe (Fig. 8). Likewise, within the G. cynanchoides complex (Fig. 7, gel panel 3) some plants hybridize to one probe but not the other, while some hybridize to JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 265 both (Fig. 8). Plants that hybridized just to the 1900 bp probe all have morphology typical of G. cynan- choides, and all bear the cpDNA haplotype S, which is the most common cpDNA haplotype with- in section Gaudichaudia, the G. cynanchoides com- plex (Fig. 3). Two of the three plants binding the 1150 bp probe also bear cpDNA haplotype S. The other plant bears cpDNA haplotype L, the most common cpDNA haplotype the G. albida complex. The samara morphology of that plant is, however, more typical of G. implexa, the amphiploids be- tween G. albida and G. cynanchoides (section Tri- tomochaudia). Most of the plants hybridizing to both probes show the samara morphology typical G. implexa. Eight out of fifteen plants binding both probes have cpDNA haplotypes from section Tri- tomopterys, indicating their affinity with the G. al- bida complex, six have cpDNA haplotypes from section Gaudichaudia, indicating their affinity with the G. cynanchoides complex, and one has a cp- DNA haplotype identified as a member of section Archaeopterys, an outgroup to section Tritomopter- ys. Experiments using primer sequence CAGGCC- CTTC (primer A-1) yielded similar results (Fig. 9). Blots were probed with two labeled fragments from the gels used to produce the autoradiographs. The first probe was made with a 1400 bp fragment col- lected from lane 47 (Jessup 4039, G. galeottiana, Puebla). The second probe was made with a 1000 bp fragment collected from lane 16 (Jessup 4055, G. albida, Chiapas). The first probe (from G. gal- eottiana) hybridized strongly to five collections from section Zygopterys, including the source lane (G. galeottiana) three lanes representing G. zyg- optera and one lane representing G. intermixteca. Lanes representing G. krusei (section Archaeopter- ys) and G. hirtella (section Oligopterys, formerly in Aspicarpa) were also hybridized by the probe. Hybridization signal from lanes representing G. krusei and G. hirtella were weak, but stronger than hybridization to lanes representing other elements of Gaudichaudia. The second probe (from G. al- bida) hybridized strongly to plants in the G. albida complex, and to several of the amphiploids. Two plants were hybridized by both probes. One of those plants (Jessup 4047, Oaxaca) carries a cp- DNA haplotype (A) from section 7Jritomopterys, the clade associated with the G. albida complex. The other plant (Anderson 12990, Oaxaca) carries a cpDNA haplotype (X) from section Zygopterys, the clade associated with the G. galeottiana com- plex. From this we can infer that Anderson 12990, carrying RAPD sites from both G. albida and G. galeottiana and a cpDNA haplotype from the G. galeottiana complex, must be derived from a cross between a plant in the G. albida complex, the pol- len donor, and a plant in the G. galeottiana com- plex, the maternal parent (Fig. 9). The hybrid, Jes- sup 4047, must have formed the other way around. The pollen donor was from the G. galeottiana com- 266 MADRONO [Vol. 49 TABLE 3. LABELED RAPD PROBES HyYBRIDIZED TO BLOTS. Numbers of bands hybridizing probes is recorded for all lanes with a strong hybridization to the probe. Lanes that were blank on the gels have been omitted. Collections are grouped by taxonomic assignment based on morphology. Asterisks indicate probe sources. Primer Probe Collec- Al Al A-2 A-3 A-7 A-19 Al9 A-19 A-19 tion # cpDNA Lane # 16:1000 47:1400 8:1250 28:950 4:850 12:14000 12:1100 75:1150 75:1900 Section Tritomoptetys Chi4056 cpDNA 1-F 1 4 2 Gro4067 cpDNA 1-D 3 Zz 4 2) 4 3 Mex13275 cpDNA 1-D 4 3 4 £2, 4 3 Oax4041 cpDNA 1-B ) 3 4 D Oax4052 cpDNA 1-L 6 3 2) 2 D 3 Gro4066 cpDNA 1-D 8 3 *A 2D 4 3 Mic13309 cpDNA 1-L 10 3 2 Mor12937 cpDNA 1-D IZ 4 *A #8) Nay4088 = cpDNA I-L 13 4 Dy) 3 S$in4102 cpDNA 1-N 14 3 4 2 3 Chi4055 cpDNA 1-C 16 oS) 4 2 2 Chil3244 cpDNA 1-K 18 It 4 D) Chil3225 cpDNA 1-K 19 3) 4 2 D) Jal4084 cpDNA 1-L aM D) Jal4085 cpDNA 1-L 22 2) Oax405 1 cpDNA 1-L 24 3 4 2 Oax13216 cpDNA 1-K 26 3 4 2 2 Chi4058 cpDNA 1-B Di 2) 4 Chi4060 cpDNA 1-B 28 4 oe Chi4061 cpDNA 1-B 29 4 2 2 4 Oax4054 cpDNA I-K 30 4 2 Oax4048 cpDNA 1-D 37) 4 4 Natural and Artificial Hybrids Zac4 106 cpDNA 1- 84 4 Zacl2624 cpDNA 2- 85 4 2 3 3} 4 Gua4006 cpDNA I- 86 4 2 3 1-L 2-S 1-L Gua4108 cpDNA 1-L Hid32522 cpDNA 1-L 88 1-L 1-L 1-L 1-L (ee) ~ iW) OO 4 2 2) 3 Tam4001 cpDNA 1- 91 4 D 1 3 4 Tam4113. cpDNA I- 92 4 2, 3 Tam4114 cpDNA 1- 93 4 2 1 4 Nue4115 cpDNA 1- 94 4 2) 1 3) 4 SLP4002. cpDNA 1-L 95 1 3 4 SLP4043, + =cpDNA 4-AD 97 1 4 2 1 3) 4 SLP4056 cpDNA 1-L 98 4 2 1 3 4 Mex4020 cpDNA-1B 102 4 2 3 Mex4021 cpDNA 1-AB 103 4 3 Mex4023 cpDNA 1-AB 104 4 2 4 Mex4025 cpDNA 1-AB 106 4 Mex4026 cpDNA 5-AE 107 4 3 Mic4074 cpDNA 1-D 108 4 2 3 SLP13316 cpDNA 1-L 109 4 D l 3 2 Jal4076 cpDNA 2-S 11] 4 3 3 Jal4079 cpDNA 2-S 112 4 2 3 5 4 Jal4080 cpDNA 2-S 3 2, 2 3 Jal408 1 cpDNA 2-S 114 2 Jal4082 cpDNA 2-S 115 2 3 Jal3707 cpDNA 1-M 116 2 I 3 + Mic403 1 cpDNA 1-L 118 3 Mic4032 cpDNA 1-L tS 4 Ss) Mic4034 cpDNA 1-L I 2 2 | Mic4035 cpDNA 2-P 122 D Mic4037 cpDNA 2-Q 124 3 4 2 2 3 Mic13291 cpDNA 1-D QS 4 2 2: Oax4043 cpDNA 4-AD 2 3 4 2 Oax4044. cpDNA 4-V 128 3 4 4 Z 3 2002] JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 267 TABLE 3. CONTINUED. Primer Probe Collec- Al Al A-2 A-3 A-7 A-19 Al19 A-19 A-19 tion # cpDNA Lane # 16:1000 47:1400 8:1250 28:950 4:850 12:14000 12:1100 75:1150 75:1900 Oax4046 cpDNA 1-A 130 p) Oax4047 cpDNA 1-A 131 2 + 2 Oax13031 cpDNA 1-A 132 4 p) H687 cpDNA 1-D 39 4 2 4 3 H2187 cpDNA 1-L 36 4 2 H2487 cpDNA 1-M 64 4 3 3 3 4 H5487 cpDNA 1-L 65 4 2 2 l 3 4 H289 cpDNA 1-L 66 4 j 3 4 H489 cpDNA 1-L 67 4 l 3 4 H4387 cpDNA 1-L 99 4 2 l 3 4 H689 cpDNA 1-L 100 4 ] 3 H4687 cpDNA 1-L 101 4 2 ] 3) 4 H2586 cpDNA 1-L 133 3 + 5) 3 H2687 cpDNA 1-M 134 3 2 3 4 Section Guadichaudia cpDNA Zac4103 cpDNA IO-G 68 ] 3 4 Zac4104 cpDNA 2-R 69 ] 3 Gua4107 cpDNA 2-R 70 l 3 3 Gua4007 cpDNA 2-R a 3 Hid4109 cpDNA 2-S 72 4 1 3 4 Hid4110 cpDNA 2-S a3 + ] 3 3 Hid4111 cpDNA 2-S 74 + 2 3 3 Hid4112 cpDNA 2-U iS + ] 23) *4 SLP4004. cpDNA 2-S 76 4 3 Jal4077 cpDNA 2-S 78 2 l 3 4 Jal4078 cpDNA 2-S TS 3 Mex82260 cpDNA 2-S 80 2, 3 Mic4075 cpDNA 2-S 81 2 ] 3 4 Mic13296 cpDNA 2-S 82 2 3 Zac4105 cpDNA 2-S 83 4 2 3 3 3 Sections Cyclopterys, Zygopterys, Oligopterys and outgroups Gro4510 cpDNA 3-AF 39 4 2 2 3 Jal4083 cpDNA 3-AB 40 = Col12699 cpDNA 3-AC 4] 4 Oax4293 =ocpDNA 1-L 44 3 4 2 l 3 4 Pue4040 cpDNA 1-A 45 4 2 Pue4039 cpDNA 4-AG 47 *4 Oax4049, = =cpDNA 4-W 49 p 4 2 3 Oax12990 cpDNA 4-X 50 3 4 4 2 Oax13138 cpDNA 4-W 51 4 2 3 Gro4069 cpDNA IO-J 54 2 Asp525 cpDNA ASP 56 2 3 Asp526 cpDNA ASP a7 2 Jan1254 cpDNA JAN 59 2 Jan3373 cpDNA JAN 61 2 plex and the maternal lineage from the G. albida complex (Fig. 9). Blots made from gels of RAPD produced with primer sequence TGCCGAGCTG (A-2) were probed with a labeled 1250 bp fragment from G. diandra (lane 8, Jessup 4066, Guerrero) (Table 3). The probe bound strongly to RAPD fragments from plants throughout the genus. About 77% of the plants sampled in the G. albida complex hybridized the probe, but only 40% of the plants sampled in the G. cynanchoides complex hybridized the probe. About 76% of the plants sampled from among the amphiploids hybridized the probe. Representatives from G. cycloptera, G. mcvaughii, and G. galeot- tiana also bound the probe. Blots made from gels of RAPD fragments pro- duced with primer sequence AGTCAGCCAC (A- 3) were probed with a labeled 950 bp fragment am- plified from an unpublished species in the G. albida complex (lane 28, Jessup 4060, Chiapas) (Table 3). About 86% of the plants sampled in the G. albida complex hybridized the probe. Only 13% of the plants sampled in the G. cynanchoides complex hy- bridized the probe. About 53% of plants sampled 268 MADRONO [Vol. 49 RAPD A-19: Primer sequence: *>CAAACGTCGG?* Probe source: lane 12, ~1400 bp; Morelos r ton 2 Fic. 5. ~N Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1) probed with 1400 bp fragment from lane 12 (Anderson 12937, G. diandra, Morelos), indicated by arrow. Fruits are illustrated in lanes showing strong hybridization to the probe for all plants with fruits available. in the amphiploid complex hybridized the probe. The probe also hybridized to one plant in the G. cycloptera complex, and two plants from the G. galeottiana complex. Blots made from gels of RAPD fragments pro- duced with primer sequence GAAACGGGTG (A- 7) were probed with a labeled 850 bp fragment from G. albida sensu stricto (lane 4, Anderson 13275, Mexico) (Table 1). The probe bound strong- ly to diverse elements of Gaudichaudia and to both collections of Janusia, another closely related ge- nus, included in the sample, but did not hybridize to G. hirtella (formerly in Aspicarpa), and was not prevalent within any of the groups sampled. About 22% of the plants sampled in the G. albida complex hybridized the probe, and about 27% of plants sam- pled in the G. cynanchoides complex hybridized the probe. Among members of the amphiploid com- plex, about 26% hybridized the probe. Two plants representing the G. galeottiana complex, and one member of the G. cycloptera complex also hybrid- ized the probe. Comparing the distribution of hybridization sig- nal across probes, two probes hybridized specimens predominantly in the G. cynanchoides complex (A- 19 75:1150 and A-19 75:1900) but hybridized very few specimens in the G. albida complex. Three probes (A-1 16:1000, A-3 28:950 and A-19 12: 1400) hybridized specimens predominantly in the G. albida complex but very little or not at all in the G. cynanchoides complex. All five of those probes hybridized samples prominently in the am- phiploid complex. Among the amphiploids, probes hybridizing predominantly in the G. albida com- plex were combined in some combination with probes hybridizing predominantly in the G. cynan- choides complex in about 26% of the plants sam- pled. DISCUSSION The assumption of homology. One of the key as- sumptions allowing the use of RAPD markers as characters is that bands occurring in different sam- ple lanes at the same position, i.e., bands having the same molecular weight, have DNA sequences sharing sequence homology, and are therefore re- lated by ancestry. One way to test that assumption JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 269 2002] RAPD A-19: Primer sequence: *>CAAACGTCGG* Probe source: lane 75, ~1150bp; Hidalgo Fic. 6. fragment from lane 75 (Jessup 4112, G. cynanchoides, Hidalgo), indicated by arrow. Fruits are illustrated in lanes showing strong hybridization to the probe for all plants with fruits available. would be to sequence several sites having the same molecular weight, but that approach is expensive and would be limited to relatively few specimens. Use of RAPD hybridization blots permits screening a large set of DNA’s, in this study representing 75 plants in the genus. The procedure probes a radio- labeled RAPD fragment of known size and source against Southern blots made from the RAPD gels. RAPD fragments on the blot hybridizing to the la- beled probe must have substantial sequence ho- mology for strong hybridization. In the experiments reported here only lanes that showed a strong hy- bridization signal were included in the data matrix. Many lanes showed weak binding of the probe and might have some sequence homology, but diver- gence was sufficient to weaken the signal. Weak hybridization is expected from the primer sequence alone. The assumption of independence. In addition to the question of sequence homology of RAPD frag- ments of the same weight across samples, there is the question of sequence homology of RAPD frag- ments of different weight within a sample. Since both the theory of how RAPD markers behave in amphiploids and the empirical evidence presented here suggests that hybrids combine distinct RAPD sites of the parental lineages, we expect that some of those sites will be homologous, or more specif- ically, synologous (Mindell and Meyer 2001 )—di- vergent and descended from a common ancestor but residing in the same genome by virtue of retic- ulate ancestry. Synologous fragments would have sufficient sequence divergence (insertions, dele- tions, substitutions) to express different electropho- retic mobility, but would retain enough sequence homology to hybridize to a probe from the synol- ogous locus. Changes in size of a RAPD site caused by insertion/deletion events are likely to develop in reproductively isolated lineages. When the lineages bearing the divergent sites merge in an amphiploid each will be expressed, resulting in complementa- tion. In the absence of recombination between pa- rental genomes the hybrid lineage would then be a fixed heterozygote. Detection of fixed heterozygos- ity corroborates other evidence supporting an am- phiploid origin of the lineage. It is tempting to in- 270 MADRONO [Vol. 49 RAPD A-19: Primer sequence: °>CAAACGTCGG* Probe source: lane 75, ~1900bp; Hidalgo 126 128 130 132 1 119 121 123%y Fic. 7. Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1) probed with 1900 bp fragment from lane 75 (Jessup 4112, G. cynanchoides, Hidalgo), indicated by arrow. Fruits are illustrated in lanes showing strong hybridization to the probe for all plants with fruits available. terpret multiple bands with sequence homology as indicating successive layers of fixed heterozygosity built up in the tiered genomes of an ancient poly- ploid complex. The evidence presented here is consistent with synologous origins for fragments that bind the probe but which have different molecular weights from that of the probe. Competing hypotheses can not, however, be ruled out with the available data. For example, multiple fragments hybridizing the probe within a specimen could indicate multiple nested priming sites within the amplification win- dow of PCR conditions used. Some PCR products would encompass three pairs of priming sites, some two, and some only one, resulting in three frag- ments sharing overlapping sequence identity. Other scenarios explaining the appearance of different sized RAPD fragments with sequence homology can be envisioned. Without a detailed study of how the coamplifying fragments are arranged on the chromosomes it is not possible to support or reject the alternative hypotheses. Introgression. A plausible explanation for the observation that all plants binding both the A-19 1900 bp and A-19 1150 bp probes show morphol- ogy typical of hybrids between G. cynanchoides and G. albida (Fig. 8) is that cryptic sibling species within G. cynanchoides are forming tetraploids and crossing, and those polyploids are capable of form- ing amphiploids with similar lineages from the G. albida complex. It is quite possible that some lin- eages in the G. cynanchoides complex (and in other groups as well) are geographically restricted where- as the polyploid lineages involved in most of the wide crosses are weedy and wide ranging and carry more of the genetic diversity as fixed heterozygos- ity. If we compare the autoradiograph of the A-19 1150 bp probe (Jessup 4112, G. cynanchoides, Hi- dalgo, lane 75), and that of the A-19 1100 bp probe (Anderson 12937, G. diandra, Morelos, lane 12) for just eight plants from the G. cynanchoides com- plex (lanes 81—88) (Fig. 10), we observe one plant hybridizing only to the 1100 bp probe (probe from 2002] Probe source: lane 75; Hidalgo mat68 70 72 #7 7% 7 79 8 1900 bp = —- 2 a> ae we oo 1150 bp Ea vie 2 eee 1900 bp blot ii (cpDNA from section Tritomopterys) (cpDNa from section Gaudichaudia) os ‘s M3 é 3 \ ; > cpDNA from id \ section Archeopterys G. cynanchoides Fic. 8. : 2 Plants hybridizing on cpDNA from section Gandichaudia | G. implexa Plants hybridizing both 1900 bp and 1150 bp probes displayed in order of appearance on blot. JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 271 RAPD A-19: Primer sequence: °>CAAACGTCGG? & ~ a. , “sy F 3 a , ly the i” 1900 bp probe; all section Gaudichaudia - s & @ 7 (cpDNA from "section Gaudichaudis) Ap 3 ne intermixteca vial saat \ cpDNA from section Tritomopterys Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1), lanes 68-101. Gel, at top, is compared with autoradiographs from 1900 bp probe and 1150 bp probe. Both probes were prepared from fragments in lane 75 (Jessup 4112, G. cynanchoides, Hidalgo), indicated by arrow. Fruits illustrated top panel are from plants hybridizing only the 1900 bp probe; fruits illustrated middle panel are from plants hybridizing only the 1150 bp probe; fruits illustrated bottom panel are from plants hybridizing both the 1900 bp and 1150 bp probes. In bottom panel sectional affiliation of cpDNA haplotypes is indicated for each fruit illustrated, and dotted lines separate described species. G. albida sens. lat.), two plants hybridizing only to the 1150 bp probe (probe from G. cynanchoides), and three plants hybridizing to both probes. The plants hybridizing both probes all carry cpDNA haplotypes characteristic of section Gaudichaudia. Two of the three plants hybridizing both probes are G. implexa (nothosection Tritomochaudia) and ex- hibit the samara morphology of the amphiploids. The plant hybridizing only the 1100 bp probe car- ries a cpDNA haplotype from section Tritomopter- ys, the G. albida complex. Of the two plants hy- bridizing only the 1150 bp probe, one carries a cpDNA haplotype from section Tritomopterys, and one carries a section Gaudichaudia cpDNA hap- lotype. One explanation for sites shared in this way is that introgression is occurring between the G. dian- dra lineages and the G. cynanchoides lineages. The 1100 bp probe (from G. diandra) hybridizes to sev- eral plants in the G. albida group, and to almost every plant in the G. cynanchoides group. Among the plants in the G. cynanchoides group, plants that exhibit the typical cynanchoid samaras hybridize only a single fragment, the 1100 bp fragment. In ie RAPD A-1: primer sequence: *CAGGCCCTTC* Probe source: lane 47, 1400 bp, Puebla G. galeottiana (section Zygopterys) G. intermixteca Oaxaca i G. zygoptera (nothosection se G. zygoptera (cpDNA from section Zygopterys) Pollen source = Pollen source = G. albida sens. lat. G. galeottiana Two plants bound both 1000 and 1400 bp probes. (_pDNA from section Tritomopterys) Fic. 9. Autoradiographs of RAPD products from primer A-1 probed (upper panel) with 1400 bp fragment from lane 47 (Jessup 4039, G. galeottiana, Puebla), indicated by arrow. Lower panel shows the two lanes with a strong hybridization to both the 1000 bp probe from lane 16 (Jes- sup 4055, G. albida sensu lato, Chiapas) and the 1400 bp probe from lane 47. the amphiploids the probe hybridizes to two or three fragments. The single G. cynanchoides frag- ment is evident in lane 81, Fig. 10 (lanes other than 81—88 are not illustrated for the 1100 bp probe). That observation is consistent with the hypothesis that G. cynanchoides, which carries a single variant (1100 bp in length), is introgressing with G. dian- dra. It is possible, though, that the probe source (Anderson 12937, G. diandra, Morelos, lane 12) is itself a fixed heterozygote. The probe hybridizes to all three fragments in the probe source lane (not illustrated). Though the evidence is suggestive, it MADRONO [Vol. 49 RAPD A-19: Primer sequence: *CAAACGTCGG* Probe source: lane 12: 1100bp fragment ~% section Tritomopterys) (cpDNA fom section Tritomopterys) "85 86 87 88 Probe source: lane 75: 1150bp fae Gh (section Gaudichaudia) (cpDNA from section jPrajemliae) Co » (cpDNA from section Gaudichaudia) Plants binding only 1150 EB probs G. implexa G. cynanchoides Plants binding both 1100 and 1150 bp probes. Fig. 10. Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1), lanes 81-88. Upper panel shows hybridization to the 1100 bp probe from lane 12 (Anderson 12937, G. diandra, Morelos). Middle panel shows hybridization to 1150 bp probe from lane 75 (Jessup 4112, G. cynanchoides, Hidalgo). Fruits illustrated in bottom panel for plants hybridizing both probes. may not be possible with the sample in hand to eliminate the hypothesis that what we are seeing 1s a shared polymorphism. Sequencing the RAPD fragments would show how length variants differ. Examining variation of RAPD fragment patterns within and among populations of each species would be a fruitful approach to the population level dynamics of introgression. Conclusion. What we can say for certain is that RAPD fragments of the same molecular weight can be reliably considered homologous sequences. We can also conclude that fragments of different mo- lecular weight within a lane, i.e., coming from the same nucleus, often represent size variants of a sin- 2002] gle RAPD site. Hybrids exhibit complementation of sites from parental lineages that are presumably fixed for different size fragments. This allows us to use the presence of fragments on the gels as binary characters in phenetic analysis of relatedness among the collections, even though different size fragments have homologous sequences. We can get even more specific in identifying the parental lineages for a given hybrid by combining information from the Southern transfer hybridiza- tion experiment with information from analysis of cpDNA restriction site data. The experiments using primer sequence CAGGCCCTTC (primer A-1) demonstrate this application of RAPD hybridization (Fig. 9). From the forgoing evidence it seems likely that RAPD fragments of different molecular weights but similar sequence, as demonstrated by strength of probe hybridization, can be used to characterize genomes within polyploids. Markers thus developed can be used to resolve reticulate an- cestry in amphiploid complexes that are otherwise intractable. ACKNOWLEDGMENTS The author thanks William R. Anderson, who gener- ously shared his knowledge and research collections and provided guidance and assistance with field work, and E. Pichersky and R. Fogel, who provided laboratory space and equipment. This research was funded in part by NSF grant BSR-8700340 to W. R. Anderson, and by NSF Doc- toral Dissertation Improvement grant BSR-8823076 to W. R. Anderson for S. L. Jessup. LITERATURE CITED ANDERSON, W. R. 1980. Cryptic self-fertilization in the Malpighiaceae. Science 207:892-893. . 1993. Chromosome numbers of neotropical Mal- pighiaceae. Contributions from the University of Michigan Herbarium 19:341—354. ARNOLD, M. L. AND S. K. Ems. 1998. Molecular markers, gene flow, and natural selection. Chapter 15 in D. E. Soltis, P. S. Soltis, and J. J. Doyle (eds.), Molecular systematics of plants Il: DNA sequencing. Kluwer, Boston, MA. Davis, C. C., W. R. ANDERSON, AND M. J. DONOGHUE. 2001. Phylogeny of Malpighiaceaea: evidence from chloroplast NDHF and TRNL-F nucleotide sequenc- es. American Journal of Botany 88:1830—1846. JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 273 DOWLING, T. E., C. Moritz, J. D. PALMER, AND L. H. RIE- SEBERG. 1996. Nucleic acids III: analysis of fragments and restriction sites. Pp. 249-320 in D. M. Hillis, C. Moritz, and B. K. Mable (eds.), Molecular system- atics, 2nd ed. Sinauer, Sunderland, MA. FEINBERG A. P. AND B. VOGELSTEIN. 1983. A technique for radiolabeling DNA restrction endonuclease fragments to high specific activity. Annals of Biochemistry 132: 6-13. FEINBERG A. P. AND B. 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Autopolyploids. Chapter 8 in First course in population genetics. Boxwood Press, Pacific Grove, CA. Lyncu, M. 1988. Estimation of relatedness by DNA fin- gerprinting. Molecular Biology and Evolution 5:384— S22). MINDELL, D. P. AND A. MEYER 2001. Homology evolving. Trends in Ecology and Evolution 16:434—440. RIESEBERG, L. H. 1996. Homology among RAPD frag- ments in interspecific comparisons. Molecular Ecol- ogy 5:99-105. SAMBROOK, J., E. E FRITSCH, AND T. MANIATIS. 1989. Mo- lecular cloning: a laboratory manual, 2nd ed. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY. WoLFE, A. D. AND A. LISTON. 1998. Contributions of PCR-based methods to plant systematics and evolu- tionary biology. Chapter 2 in D. E. Soltis, P. S. Soltis and J. J. Doyle (eds.), Molecular systematics of plants Il: DNA sequencing. Kluwer, Boston, MA. MADRONO, Vol. 49, No. 4, pp. 274-284, 2002 LONG-TERM POPULATION DYNAMICS OF NATIVE NASSELLA (POACEAE) BUNCHGRASSES IN CENTRAL CALIFORNIA JASON G. HAMILTON! Department of Ecology, Evolution and Marine Biology, University of California, Santa Barbara, CA 93106 JAMES R. GRIFFIN AND MARK R. STROMBERG2 Hastings Natural History Reservation, Museum of Vertebrate Zoology, University of California, 38601 E. Carmel Valley Road, Carmel Valley, CA 93924 ABSTRACT California bunchgrass communities are one of the most endangered ecosystem types in the United States. In this study, we sought to determine long-term (52+ years) changes in populations of native bunchgrasses, Nassella pulchra (A. Hitchce.) Barkworth and Nassella cernua (Stebb. & Love) Barkworth, in unmanaged stands. At the landscape scale, Nassella has increased. However, population dynamics of individual stands appeared related to land-use history. Non-native annuals, by themselves, did not seem to cause decline of Nassella stands, but light grazing did cause reduction of Nassella basal cover. Areas that were historically cultivated supported Nassella stands with lower basal cover and size distributions qualitatively different from areas that were never cultivated. Mortality of Nassella was concentrated in small plants. Interspecific interference probably was important in limiting seedling recruitment in stands with low Nassella basal cover, and intraspecific interference appeared to become important as Nassella basal cover increased. Even in the presence of non-native annuals, Nassella stands in areas that have not been disturbed by cultivation do not appear to require management for maintenance. New individuals are recruiting into populations, and conservative a estimate of longevity of large individuals of Nassella is 100 years. However, in areas that have been cultivated, active management may be required to increase the abundance of Nassella. Key Words: California, Nassella, grassland, survival, longevity, long-term. INTRODUCTION Perennial bunchgrass communities are one of the rarest plant communities in California (Keeley 1989, 1993) and are considered to be one of the most endangered ecosystem types in the United States (Noss et al. 1995; Peters and Noss 1995). Since the founding of the Spanish missions in the mid-1700’s, massive invasions of annual grasses from the Mediterranean basin have altered native communities to such a degree that today the origi- nal extent and composition of these communities is unknown (Keeley 1989; Heady et al. 1992; Ham- ilton 1998). Today in California, an area of approx- imately 7,000,000 ha is dominated by non-native annual grasses (Huenneke 1989). In many cases, these non-natives comprise from 80% to 100% of the cover (Biswell 1956; Heady 1956; Macdonald et al. 1988; Heady et al. 1992), and the small patch- es of perennial bunchgrasses that still exist in Cal- ifornia (including Nassella (=Stipa) pulchra (A. Hitche.) Barkworth and closely related Nassella cernua (Stebb. & Love) Barkworth) always include non-native grasses. ' Present address: Biology Department, 161 CNS, Ith- aca College, Ithaca, NY 14850-7278. * Author for correspondence, e-mail: socrates.berkeley.edu stromber @ In most of California, the original community composition of areas in which Nassella bunchgrass- es are found today is a matter of conjecture (Ham- ilton 1998). However, it is clear that over the past two hundred years, the biotic environment has changed dramatically for these bunchgrasses (Dyer and Rice 1999). There are no ‘pristine’ areas of California grassland left. Non-native annual grasses such as Bromus hordeaceous L., Bromus diandrus Roth, Avena fatua L., and Avena barbata Link have invaded every known bunchgrass stand. Further- more, due to land clearing, farming, and extreme over-grazing (Burcham 1957), even areas that are currently protected have been previously disturbed in some manner. Because of a lack of long-term studies, it is unknown whether a new steady-state situation has been achieved in the California grass- lands, or whether bunchgrass stands are still ad- justing to the altered conditions. In California, there is a growing interest in res- toration and conservation of Nassella bunchgrass communities (Knapp and Rice 1994; Stromberg and Kephart 1996; Carlsen et al. 2000; Kephart 2001). Attempts at generalized management pre- scriptions that promote grazing and/or burning as a tool to reduce competition from annual grasses and enhance longevity of mature bunchgrasses have been proposed (e.g., Menke 1992) for inland sites. Substantial differences are evident between inland 2002] and coastal native grasslands where Nassella is a co-dominant (Stromberg et al. 2001). On inland sites, recruitment of Nassella appears to be limited by competition by non-native annuals (Dyer and Rice 1999) and management strategies have been developed to improve establishment by reducing exotic seed banks (Stromberg et al. 2002). Studies of mortality and recruitment, along with restoration, are lacking in coastal environments of California and management strategies suggested or inland sites (Menke 1992) may require modification. We lack the fundamental information concerning long- term stability characteristics of Nassella stands in the face of competition from non-native annual grasses. Because of this, land managers have been forced to rely on hearsay to determine whether na- tive grasslands require management in order to per- sist, and if so, what kind. Results from short-term studies (one-two years) have tended to be unreli- able indicators of longer-term dynamics. For ex- ample, in one study, preliminary results after 16 months indicated that burning and early-spring grazing were effective at increasing Nassella pul- chra seedling establishment and survival (Fossum 1990). However, in the same study, after four years, it was concluded that burning and grazing were not effective at enhancing Nassella pulchra seedling re- cruitment (Dyer et al. 1996). In our study, we sought to determine long-term changes in populations of Nassella bunchgrasses in unmanaged stands. In particular, we asked: (1) In the absence of fire or grazing, has Nassella in- creased or decreased at the landscape scale? (2) At the scale of individual stands, are established pop- ulations of Nassella stable? (3) At the scale of sin- gle individuals, is there life-stage-related mortality that suggests interference from non-native grasses? (4) Do trends in multi-scale population dynamics suggest that Nassella requires management for per- sistence when there is interference from non-native annuals? There are very few sites in California where data exist that allow for analysis of long-term trends in Nassella bunchgrass populations. One such place is the Hastings Natural History Reservation in the foothills of the South Coast Range in central Cali- fornia. Here, in a study initiated in 1944 by G. L. Stebbins, White (1966) described old field succes- sion Over a 22-year period. Using unpublished data from studies by both Stebbins and White and more detailed data from a number of other bunchgrass stands at Hastings Reservation, we have been able to extend the original findings of White to encom- pass a period of 52 years, and to compare a number of sites around Hastings Reservation representing many ecologically distinct situations with different land-use histories. METHODS The 911-ha Hastings Natural History Reserva- tion (36°33'30"’N, 121°33'30”W) is located in the HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 275 interior foothills of the coastal Santa Lucia Moun- tains in central California, 33 km southeast of Mon- terey. The Mediterranean climate, characterized by hot, dry summers and cool wet winters, supports a number of plant communities including oak wood- land, chaparral, and grassland (Griffin 1971; MacRoberts and MacRoberts 1976; Williams and Koenig 1980). Hastings Reservation has been pro- tected from fire and grazing since its establishment in the fall of 1937, except for a 40 ha horse pasture that was lightly grazed until 1968. Between 1944 and 1977, several plots were es- tablished at sites with a variety of land use histories (all within four km of each other) around Hastings Reservation to monitor native Nassella (including both N. pulchra and N. cernua) bunchgrasses (Ta- ble 1; detailed maps showing stand locations are available at Hastings). Original plots were estab- lished to monitor small patches of Nassella that re- mained, for whatever reason, in what was often a much larger expanse of introduced, annual grasses on abandoned fields or in oak savanna. Because data collection for these plots was not coordinated, available data differed for each plot. Data collected ranged from multiple censuses of size and location of every Nassella individual in a given plot, to sin- gle censuses indicating only presence or absence in a plot. The two species of Nassella that occur at Hastings Reservation are extremely similar in veg- etative morphology and are known to hybridize (Stebbins and Love 1941; Love 1954). Because many of the data sets for plots did not differentiate between these closely related species, we did not differentiate these species in our data analysis. All nomenclature follows Hickman (1993). We used data from nine plots (Table 1) that were established in four areas at Hastings Reservation: White Prairie (one plot), South Sandstone (one plot), North Field (five plots), and Arnold Field (two plots). White Prairie, South Sandstone, and North Field are all within a few hundred meters of each other (elevation ca. 550 m), and Arnold Field (elevation ca. 730 m) is located approximately four km SW of the other three areas. Nassella bunch- grasses (also known as tussock grasses or tufted grasses) are perennial grasses that have a clumped or cespitose growth habit. For all plots, Nassella individuals were defined as any physically distinct tussock that was not clearly a clonal fragment from some larger tussock (see e.g., Wilhalm 1995). An analysis of size structure of the Nassella pop- ulations was based on historical data or 1996 mea- surements of basal diameter measurements of in- dividuals. For many plots, historical data were tak- en from detailed tracings or maps of individual plants. Plants were divided into size classes based on basal diameters: (1) less than or equal to one cm, (2) greater than one cm to 5 cm, (3) greater than five cm to 10 cm, and (4) greater than 10 cm. Plants that were not circular were assigned to di- ameter classes based on the corresponding basal ar- 276 MADRONO [Vol. 49 TABLE |. SUMMARY INFORMATION FOR PLOTS USED IN THIS STUDY. Notes Plot area Date Plot name (m7) established White Prairie 10 1977 South Sandstone 10 1966 North Field 427 149 1966 North Field 428 56 1966 North Field 429 84 1966 North Field 409 9 1965 North Field 412 9 1964 Arnold 420 9 1964 Arnold 449 10 1979 Considered to be undisturbed relict of bunchgrass prairie; moderate gopher activity Never cultivated; in a 40 ha area lightly grazed by 2—5 horses 1940-1968 Originally woodland; lightly cultivated for barley ca. 1860— 1937 Originally woodland; lightly cultivated for barley ca. 1860— O37 Originally blue oak woodland; lightly cultivated for barley ca. 1860-1937 Sub-plot of North Field 429 Originally valley oak savanna; cleared; cultivated as vineyard ca. 1920-1937 Originally valley oak savanna; cleared; cultivated barley ca. 1860-1937; many gophers Probably lightly cultivated 1860—1937; burned in 1979, many gophers eas. To estimate minimum longevity of Nassella in- dividuals, we used direct tagging of plants. We es- timated the age of large individuals by determining average rates of increase in basal area and calcu- lating the number of years required for an individ- ual to attain a given size. The plot in White Prairie was established in 1977. This plot is surrounded by oak woodland and, as there is no record of land clearing, the original vegetation was probably grassland. White Prairie is considered to be a relict of pre-European Nassella bunchgrass grassland because it has not been cleared, and was probably only occasionally grazed. The plot showed evidence of moderate go- pher (Thomomys bottae) activity. In 1977, individ- ual Nassella plants were tagged, and, in many cas- es, wire loops were placed around the base of the plants to ensure future identification of individuals. Historical data include scale maps showing location and shape of each Nassella individual, as well as basal diameter measurements. In most cases Nas- sella individuals were generally elliptical or circu- lar, and, because of the detail of the maps, it was possible to identify individuals that were more ir- regular in shape. A digital image analysis system (Decagon Devices, Pullman Washington) was used to calculate basal area of irregular-shaped clumps. In the 1993 census, many tags could not be re- located; however, carefully drawn maps from 1977 allowed us to identify most individuals. The plot in South Sandstone was established in 1976. This plot is located adjacent to oak wood- land, and, like White Prairie, the original vegetation was probably grassland. The South Sandstone plot was never cleared or cultivated, but is located in a 40-ha pasture that was lightly grazed by two to five horses until 1968. In 1976, individual Nassella plants were tagged using the same methods as pre- viously described. Historical data include scale maps showing location and shape of each Nassella individual, as well as basal diameter measurements. Almost all original tags in the South Sandstone plot were re-located in the 1993 census and, thus, tem- poral comparisons are always on the same individ- uals. A separate study using the South Sandstone plot counted Nassella seedlings in 1976, 1977, 1978, and 1979. The five plots in North Field were established between 1964 and 1966 (Table 1), and are all lo- cated within about 60 m of each other. Before 1900, North Field was probably oak woodland dominated by Quercus douglasii (White 1966). Around 1900 the trees were cleared, and the relatively level plots 427, 428, and 429 (including 409, a subplot of 429) were cultivated for barley. Plots were located on clay-sand soils with rock outcrops. Although the field was cultivated for small grain production, only mule-drawn implements were used. Isolated, rocky outcrops where Nassella persisted were relatively undisturbed by the light cultivation equipment used during farming. North Field plot 412 occurs on a slope and was a vineyard. Data for North Field plots 427, 428, 429, and 409 from 1966 (1964 for plot 409) were taken from maps on which number and approximate location (but not size) of Nassella individuals were recorded. In the 1996 re-census, in order to calculate basal cover, two orthogonal diameters were recorded for every individual and basal area calculated (assuming an ellipse). Nassel- la individuals in North Field plot 412 were mapped over a period from 1964 through 1996. In 1977, 1993, and 1996, basal area of each individual was also measured. All five North Field plots are in- cluded in a larger area that had been monitored since 1944. Available data from 1944 through 1964 indicate only whether or not Nassella bunchgrasses were present in the larger area. The two plots in Arnold Field were established 2002] HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL TABLE 2. TOTAL BASAL AREA OF NASSELLA IN PLOTS AT HASTINGS RESERVATION (cm2?/m72). White South North Year Prairie Sandstone Field 412 1964 e @ 9 1976 @ 740 ® 1977 1140 770 370 1993 1190 1180 0) 1996 e e 19 in 1964 and 1979 (Table 1). Before 1937, these two plots were probably dominated by chaparral, since they occur near stands of Adenostoma fasciculatum Hook. & Arn., and 20% of Arnold 449 was covered with this shrub in the 1996 re-census. Plots in Ar- nold field area also on relatively shallow soils with rocks near the surface, and may not have been deeply tilled with the mule-drawn discs used at the time. Both Arnold plots show evidence of very high gopher activity. Historical data include scale draw- ings of Nassella individuals in Arnold 420 (used to obtain information on density and basal diameter of Nassella in 1964) and rough drawings for Arnold 449 (used to calculate density). In the 1996 re-cen- sus, basal areas were calculated as for the other areas. RESULTS Basal area of Nassella showed different patterns of change in different plots (Table 2). The relict White Prairie plot was essentially constant over the 16-year period from 1977 to 1993. Nassella at South Sandstone increased in basal area over this TABLE 3. 277 North North North Arnold Field 427 Field 428 Field 429 420 6 * @ 400 @ e ry @ @ @ ® @ @ @ € € 250 310 290 670 time period, achieving in 1993 a value essentially the same as White Prairie. There were insufficient data to draw conclusions about changes in total bas- al cover of Nassella in North Field, except for plot 412 (former vineyard). In this case, Nassella dis- appeared between 1977 and 1993; however, in 1996, a very small amount of Nassella was again found in this plot. In Arnold 420, Nassella basal area increased between 1964 and 1996. Density of Nassella individuals also showed dif- ferent patterns of change in different plots (Table 3). The relict White Prairie plot had the highest density of all measured plots in 1977 and exhibited a slight increase with time. Mature Nassella indi- viduals at South Sandstone decreased by about 50% between 1976 and 1993, even though Nassella seedling recruitment pulses temporarily increased total Nassella density in 1978 and 1979. In North Field 412 (former vineyard), density of Nassella increased from 1951 until 1970, and then declined to zero by 1993. In 1996, three individuals were again found in this plot. In North Field plots 427 and 428, Nassella density increased between 1966 DENSITY OF NASSELLA (PLANTS/m?) FOR PLOTS AT HASTINGS RESERVATION. Missing entries indicate that plots were not yet established or were not measured in that year. * Density of mature individuals only is 12.2. ° Density of mature individuals only is 11.7. South White Sand- North North Year Prairie stone Field 412 Field 427 1944 6 @ none none 1945 ® ® none none 1946 6 e none none 1947 e e none none 1951 e ® present present 1964 ® ® 1.8 present 1965 e e 1.9 ® 1966 é e ® OF 1969 6 ® 6.1 e 1970 e e 7.8 e 1972 e e 327, e 1974 * ® 4.4 ® 1976 e 12.8 @ e 1977 13.8 i sey 5.4 ® 1978 | he 60.4 e e 1979 ® 48.0° e @ 1984 ® 6 | e 1991 e e 1 e 1993 14.7 6.9 0 ® 1996 @ e 0.3 2.9 North North North Field Arnold Arnold Field 428 Field 429 409 420 449 none present present e e none increase present e e none increase present e e none stable present e & none increase present e ® present increase 2.8 33 ® e 2.9 6 ® 0.8 1.76 e e e ® ® 3.1 @ & 6 ® 4 a « e 6 é @ @ @ e 0.89 @ e § « ® ® * @ e « « é 6 6 e e 4 ® ® e « [2:5 * ® « ® 2 Jo 6 e @ e ® e ® ® « « 3 2 2.4 12.3 6.7 278 White Prairie CI) 1977 60 1993 CI 1977 60 1993 40 20 0 > S) = o = roy = Le got North Field 412 got North Field Composite 1996 40 20 $1.0 1.0-5.0 5.0-10.0 >10.0 Diameter size class (cm) Fic. 1. Size structure of Nassella populations for differ- ent stands at Hastings Reservation. Plots in North Field were measured only once. Size structures of North Field plots 427, 428 and 429 did not differ and thus the com- posite of these three plots is shown. and 1996. In North Field plot 428, Nassella density increased between 1944 and 1966, and although density in 1966 was very similar to that in 1996, measurement of the subplot of 428 (North Field 429) indicated that there were fluctuations in den- sity over this time period. Nassella density in the two plots in Arnold Field changed in opposite di- rections. Arnold 420 had greater density in 1996 than in 1964. Density in Arnold 449 decreased from 1978 to 1996. Size structure of Nassella populations varied be- tween plots (Fig. 1). The White Prairie population showed little difference in size structure between 1977 and 1993 (Fig. 1). Numbers of individuals in the three larger size classes were approximately equal to each other, and about three times more numerous than individuals in the smallest size class. At South Sandstone, in 1977, the smallest size class and the largest size class contained roughly equal numbers of individuals, and the two intermediate size classes were only slightly less nu- MADRONO [Vol. 49 100 South Sandstone White Prairie Percent mortality of given size class S © 1.0-5.0 5.0-10.0 Diameter size class (cm) in 1977 =120 >10.0 Fic. 2. Mortality as a percent of given size class for two Nassella populations at Hastings Reservation. merous. However, in 1993, size structure of the stand had come to resemble White Prairie: the smallest size class had disappeared entirely, num- bers of plants in the one to five cm size class had decreased significantly relative to 1977, and the larger two size classes increased in number. For North Field plots, it was not possible to determine changes in population size structure because we had only one observation for each plot. In 1977, North Field 412 (former vineyard) had the most Nassella individuals in the five to 10 cm size class, with fewer larger and smaller plants. Size structure in North Field 427, 428, and 429 in 1996 was very similar to that of White Prairie in 1993. Because size distributions of the three North Field plots were very similar, only a composite of these plots is shown. In 1964, Arnold 420 had most plants in the larger two size classes, with no plants in the smallest size class and very few in the one to five cm class. In 1996, the largest two size classes had decreased significantly, the smallest size class was represented, and the one to five cm size class was the largest. Mortality patterns were very similar in the two plots (White Prairie and South Sandstone) for which individuals could be identified over time (Fig. 2). Most mortality between 1977 and 1993 was in the smallest size class, with decreasing per- centage mortality for larger size classes. At South Sandstone, total mortality in the three larger size classes combined was only eight individuals. White Prairie showed only slightly greater mortality in the three larger size classes. Seedling recruitment was sporadic in space and time (Table 4). No general pattern of recruitment connected to either yearly average rainfall or 2002] HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 279 TABLE 4. NASSELLA SEEDLING RECRUITMENT (SEEDLINGS/m*) FOR PLOTS AT HASTINGS RESERVATION. Missing entries indicate that plots were not yet established, or were not measured in that year. White South North Field Year Prairie Sandstone 412 1964 e ® 0) 1965 ® ® 0.2 1969 ® 8 40 1970 e e 1.3 1972 e e 0.2 1974 e ® 0.1 1976 ® | Remi 8 1977 0.7 0.6 0.5 1978 0.8 49.8 @ 1979 e 34.1 @ 1984 e e 0) 199] ® r) 0 1993 0.7 0.1 0) 1996 e e 0 monthly average rainfall was found. In years with seedling recruitment, seedling mortality was very high the first spring, and declined with time (Fig. 3). At South Sandstone, eight of 131 small individ- uals noted in 1976 survived to 1993. Of the six Nassella seedlings noted in 1977, one survived to 1993. Although individuals recruited after 1977 were not followed between 1978 and 1993, 24 new individuals recruited into the South Sandstone pop- ulation. At White Prairie, a comparison between maps made in 1977 and 1993 indicates 50 new in- dividuals in the population. However, some of these are likely clonal fragments of previously existing individuals. Minimum longevity measurements for Nassella were made using tagged individuals. At White Prai- rie, 38 plants tagged in 1977 could be unequivo- cally identified in 1993. Of these 38 individuals, 22 had increased in basal area over this 16-year period, 500 aq iy 1978 cohort “E 400 | ro) 1979 cohort = es < | a & 300 l = = | o ® 599 |. 1976 | S cohort Fs} | = 3 100 | | 1977 cohort a 1975 1980 1985 1990 1995 Year Fic. 3. Seedling mortality over time for four cohorts of seedlings at Hastings Reservation South Sandstone plot. North Field North Field Arnold 427, 428, 429 409 420 e 0) O 8 @) ® ad 0.2 ® e 1.6 e e ® ® @ O @ ® ® ® ® € © ® @ ® @ © ® ® ® ® 8 ® © co) ® 0.03 0.1 Lee indicating that they were still vigorous after this amount of time. At South Sandstone, 69 plants were identified as being more than 17 years old, and of these, 51 had increased in basal area. At North Field 428, 12 plants were staked in 1966 and one of these was still alive thirty years later. At Arnold 449, 18 tags were re-located after 17 years, five of which still had living Nassella individuals. Actual age of large Nassella individuals was es- timated using the average rate of increase of basal area over a 16-year period (Table 5). Due to large variation, long-term average growth rates were not significantly different between size classes or be- tween sites. The largest individuals at White Prairie and South Sandstone had basal areas between 400 and 700 cm’. Using the average long-term growth rate for both sites and all size classes (5.0 cm/?/ year), these individuals were calculated to be 80 to 140 years old. Accumulation of dead material in individual Nassella tussocks does not predict individual mor- tality. In 1976, maps of the South Sandstone plot TABLE 5. INCREASE IN BASAL AREA BY SIZE CLASS (cm?/ YEAR) OF NASSELLA IN PLOTS AT HASTINGS RESERVATION. Only plants that increased in size over time period are included. Data are averages + | standard deviation. Num- bers in parentheses are numbers of plants. Beginning White Prairie South Sandstone size class 1977 to 1993 1976 to 1993 =1 cm none S.2232 2:8 68) (<=0.78 cm?) 1 cm to 5 cm 4.2 + 2.9 (9) Sos2s5,0°(04) (0.78 to 19.6 cm?) 5 cm to 10 cm 2STEUL ST (S) 4.7 + 6.4 (14) (19.6 to 78.5 cm?) >10 cm 8.5 + 6.8 (4) Sb a.) (>78 cm?) 280 indicated 24 individuals that had large regions of dead material. In 1993, only two of these individ- uals had experienced large declines in living basal area. Between 1976 and 1993, many individuals that initially seemed to be senescing or fragmenting rebuilt their tussocks and increased in living basal area. DISCUSSION Landscape-scale Dynamics of Nassella Without Fire or Grazing In general, Nassella bunchgrasses have increased at Hastings Reservation in the absence of fire and grazing. Both our data and those of White (1966) indicate that Nassella colonized new areas from 1944 to 1966 (Table 3). As Nassella has spread to new areas, total average basal cover in established stands has remained stable. In 1967, average basal cover of Nassella for 13 protected stands at Has- tings Reservation was 10% (White 1967). For a subsample of these areas, we also found a total av- erage basal cover of 10% in 1977 (White Prairie and South Sandstone) and again in the 1990’s (White Prairie, South Sandstone, and Arnold Field). At finer resolution, we found a difference in plots that were historically cultivated and those that were not. In the 1990’s, average basal cover in formerly cultivated plots was only about 4%, compared to 12% for plots that were never cultivated. The only location where Nassella has not maintained itself is in the former vineyard. Stand-scale Dynamics of Nassella Non-native annuals are present in varying levels all these stands (Stromberg and Griffin 1996) and by themselves, did not seem to be sufficient to cause stand declines. At White Prairie, where the only known disturbance was the historical intro- duction of non-native annuals, (1971-1991 cover of non-native, annuals was 33.1%) there was essen- tially no change in basal cover, density, or size dis- tribution of Nassella over a 16-year period of this study. It is possible that one or more of these mea- sures showed a transient initial reaction to non-na- tive annuals when they first became important in the 1800’s. However, at another Nassella grassland site in California, micro-fossil evidence suggests that density, at least, has not been affected by non- natives (Bartolome et al. 1986). Although non-native annuals did not appear to impact Nassella, even light grazing did cause sig- nificant changes in Nassella stands. Over the same period that the ungrazed Nassella stand at White Prairie was stable, the stand at South Sandstone, which had been formerly grazed, increased both in total basal cover and in changes in size distribu- tions. By 1993, total Nassella basal cover at South Sandstone was indistinguishable from the relict White Prairie plot. This suggests that a basal cover MADRONO [Vol. 49 of around 1000 cm?/m* may be the maximum that can be supported in these stands at Hastings Res- ervation. Spacing of large individuals probably re- flects long-term competition for water in limiting years, and for small individuals, competition with alien annual species (Dyer and Rice 1997, 1999). Areas that were disturbed by soil cultivation sup- ported stands with lower total cover of Nassella, and had size distributions that are qualitatively dif- ferent from areas that were never cultivated. It is not clear whether cultivation was a disturbance that permanently altered the ability of areas to support Nassella stands (Stromberg and Kephart 1996), or whether recovery is simply extremely slow. How- ever, there is little indication that stands in formerly cultivated areas are developing toward patterns similar to never-cultivated plots. Again, the only plot where Nassella disappeared was the site of the former vineyard. It is possible that differences between cultivated and non-cultivated plots are due simply to pre-ex- isting site differences from before 1900. However, both the White Prairie plot and the South Sandstone plot are associated with oak woodland, as was North Field originally. Furthermore, another study that included 80 sites from around Hastings Res- ervation and the Carmel Valley also concluded that Nassella and a number of other native plant species (e.g., Poa secunda J.S. Pres] and Chlorogalum pomeridianum (DC.) Kunth) are rare in sites that have been subjected to historic cultivation (Strom- berg and Griffin 1996). Life-stage Mortality of Nassella in the Presence of Non-native Annuals We found that mortality was concentrated in the smaller size classes, and that initial seedling mor- tality was very high. Similar results were found in a study of individually marked N. pulchra in the San Juaquin valley of California (Marty 2002). Dif- ferential mortality of young plants is very common (Sarukhan et al. 1984 and references cited therein), and studies have also noted very high seedling mor- tality for Nassella (Bartolome and Gemmill 1981; Dyer et al. 1996). However, it is unclear both the degree to which interference (sensu Harper 1961; Muller 1969) is involved, and the relative impor- tance of inter- versus intraspecific interference in this mortality. In plots where density of mature Nassella individuals was low, interspecific interfer- ence from non-native annuals is probably a primary factor leading to high seedling mortality and lim- iting seedling recruitment. Studies have shown ex- plicitly that interference from non-native annuals is detrimental to performance of Nassella individuals (Nelson and Allen 1993; Dyer and Rice 1997; Hamilton et al. 1999). However, N. pulchra seed- lings can recruit into areas dominated by non-native annuals, although competition for soil moisture greatly reduced their growth (Hamilton et al. 1999). 2002] The ability of Nassella or other native, perennial grass seedlings to thrive in soil dominated by non- native annuals may be influenced by soil microbial communities. Robinson (1971) found that Nassella seeds planted in Hastings soils dominated by Avena had significantly lower survival and growth com- pared to seeds grown in soils from relict stands of Nassella. Steenwerth (2002) found that Hastings old fields, as well as nearby recently tilled fields of the similar soil, had dramatically different micro- bial communities compared to similar soils in un- disturbed, relict Nassella stands. Indeed, a simple innoculum of soil from non-native grasslands ap- pears to inhibit the growth of native California grasses (Subramaniam et al. 2001). When basal cover of mature Nassella individuals is high, intraspecific interference may be more im- portant than interspecific competition as a factor in Nassella seedling mortality. High seedling recruit- ment appeared to be associated with low basal cov- er of mature Nassella individuals (despite presence of non-native annuals) and declined as Nassella density increased. For South Sandstone in particu- lar, as total basal cover of Nassella became similar to White Prairie, seedling recruitment densities be- came similar. Intraspecific competition for soil re- sources in established bunchgrass stands in semi- arid grasslands often causes seedling recruitment to be very low in the absence of disturbance (Aguilera and Lauenroth 1993b; Hook et al. 1994; Aguilera and Lauenroth 1995). It has been suggested that the bunchgrass growth form may be inherently associated with an increas- ing risk of death as tussocks increase in size (Harp- er 1977). This could be due to accumulation of plant litter in the tussock in situations where grasses are not subjected to periodic fire or grazing. Thus, lack of fire or grazing could lead to senescence of adult Nassella individuals (Menke 1992). This does not appear to be the case for established stands at Hastings Reservation, an inland dry grassland co- dominated by Nassella. Mortality of large individ- uals in plots that were never cultivated was very low, despite the presence of non-native annuals. For example, South Sandstone showed no mortality in the largest size class and individuals that appeared to be fragmenting demonstrated the ability to re- build the tussock. Indeed, tussock fragmentation is very common in perennial bunchgrasses (Wilhalm 1995), and individual tussocks would be expected to possess the ability to rebuild through activation of dormant meristems (Bell 1984; White 1984). There was a small amount of mortality among large individuals at White Prairie, but this was probably due, at least in part, to gopher activity (Stromberg and Griffin 1996) and did not impact total Nassella cover. In contrast, there was evidence of mortality of large individuals in formerly cultivated plots. It is possible that in these areas altered site conditions reduce longevity of Nassella individuals or that in- terference from non-native annuals is more detri- HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 281 mental in these locations. Gopher activity could also increase mortality of Nassella, and, while this might be important in Arnold Field, we observed little evidence of gophers in North Field. Responses to grazing in wetter, coastal California grasslands may be dramatically different (Stromberg et al. 2001) and longevity and responses to management may vary between inland and coastal California Nassella grasslands. Contrary to some predictions (e.g., Menke 1992), tagged plants in our plots indicate that mature Nas- sella individuals are vigorous after as much as 30 years of protection from grazing and burning. Age of clonal plants, such as Nassella, is impossible to measure directly (Stebbins 1950), so use of rates of clonal spread to estimate age is commonly em- ployed (e.g., Harberd 1961; Harberd 1962). Esti- mates of longevity of Nassella based on average growth rates indicate that large individuals are like- ly more than 100 years old. Because Nassella in- dividuals break up into clonal fragments, as do many bunchgrasses (e.g., Lord 1993; Samuel and Hart 1995; Wilhalm 1995), individuals may persist in a series of clonal fragments much longer than this. In fact, clonal fragmentation is an alternative mechanism for recruitment of ‘new’ individuals into populations (Crampton 1974; Lord 1993). Al- though growth rates for individual clones can de- pend strongly on intraspecific competition (Aguil- era and Lauenroth 1993a; Dyer and Rice 1997), interspecific competition (Nelson and Allen 1993; Dyer and Rice 1997; Hamilton et al. 1999), and genotype (Samuel and Hart 1995; Skalova et al. 1997), our age estimate is not unusually high. Stud- ies of other species of bunchgrasses have also found that individuals can be very long-lived, with estimates ranging from 450 or 500 years or even longer in undisturbed areas (Coffin and Lauenroth 1988: Lord 1993; Lauenroth et al. 1994). The Requirement for Management in Nassella Stands Information on long-term population dynamics of native perennial grasses that have been left un- disturbed by humans, is necessary to develop ef- fective prescriptions for restoration and manage- ment of areas with native perennial grass stands. Large protected areas, such as Hastings Reserva- tion, provide important reference systems in which to gather such information (Bock et al. 1993). Be- cause there are no unaltered pre-European grass- land communities remaining in California, there is no naturally occurring large model by which to es- tablish goals or measure success of management or restoration efforts. Models for restoration of Cali- fornia native grasslands depend on observations of species dynamics in small patches. Such small patches may have been the common expression of native California grasslands and large homogenous stands of native grasses were rare. However, since 282 pre-European California grasslands probably con- tained many annual and perennial grasses and forbs (Heady 1977; Heady et al. 1992), one possible goal would be to promote overall diversity of native grassland species. This is complex because some management practices that could potentially benefit Nassella appear to be detrimental to other native bunchgrass species (Dennis 1989). Therefore, lack- ing detailed, species-specific information on the ef- fects of potential management practices, landscape- scale management should be conservative. Because Nassella bunchgrasses that have been protected from grazing and burning show no signs of disap- pearing at the landscape scale at Hastings Reser- vation (despite the introduction of non-native an- nuals), landscape-scale management to maintain relict Nassella in the landscape appears to be un- necessary. Stand-scale population dynamics are site depen- dent and appear to be related to land-use history; stand-scale management should take this into ac- count. At Hastings Reservation, some of the bunch- grass stands probably pre-date European arrival, while other stands occur in sites that have been converted from other vegetation types such as oak woodland or chaparral. This is true for other areas of California as well (Huenneke 1989; Keeley 1993; Hamilton 1998). In areas that have never been cultivated, interference from non-native an- nuals does not seem to cause Nassella stand de- cline. Despite high seedling mortality and sporadic seedling recruitment, areas that were never culti- vated can have high basal cover of Nassella and some achieve replacement levels of recruitment. Our longevity estimates, and the demonstrated abil- ity of Nassella to repair dead portions of a tussock, suggest that many large Nassella individuals found in areas such as White Prairie and South Sandstone could have been present at Hastings Reservation before the area was first homesteaded in 1863 (White 1967). For such long-lived species, even if conditions allowing for seedling recruitment are quite rare, stands can achieve replacement recruit- ment (Noble 1986; Lauenroth et al. 1994). Thus, lack of seedlings is not necessarily an indication of future stand decline. Nassella stands in areas that have not been disturbed by cultivation do not ap- pear to require management for maintenance. Even light grazing appears to greatly reduce Nas- sella basal cover. Although Nassella can sometimes persist in grazed areas (White 1967; Stromberg and Griffin 1996), intensive, year-round grazing on in- land stands seems to have the potential to be det- rimental to stand persistence. More studies are re- quired before decisions can be made concerning the compatibility of Nassella with grazers, or the use of grazing as a management tool as some have sug- gested (e.g., Menke 1992). Cultivation appears to be the most detrimental disturbance, and left alone, Nassella seems to re- cover extremely slowly (if at all) from a distur- MADRONO [Vol. 49 bance of this type. Other studies have come to a similar conclusion (Stromberg and Griffin 1996). Nassella stands that were historically cultivated have low basal cover and do not appear to be de- veloping toward the condition of stands that were not subjected to cultivation (e.g., Fig. 2). Further- more, Nassella in at least one population in a his- torically cultivated stand has declined significantly. At Hastings Reservation, some of the Nassella stands that were historically cultivated occur in ar- eas that have been converted to grassland from oth- er vegetation types. In such cases, the appropriate management goal may be simply to promote Nas- sella. In formerly cultivated areas, there are few large individuals, and this suggests that Nassella may be shorter-lived in these areas than in areas that were never cultivated. Low basal cover in areas subjected to historic cultivation may be the result of seedling recruitment not being frequent enough for stand replacement or growth. Therefore, in areas with historical cultivation, active management to increase seedling survivorship, while not harming mature plants, may be required to increase abun- dance of Nassella. ACKNOWLEDGMENTS The UC Hastings Reservation made this long-term work possible. We thank Thomas Cate and Laura Rosen- feld for field assistance. We are grateful to J. R. Haller, Claus Holzapfel, Bruce Mahall, Elizabeth Painter, Laura Rosenfeld, Jochen Schenk, Ed Schneider, and Josh Schi- mel for providing helpful comments on the manuscript. 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PRIGGE Herbarium—Botanical Garden, University of California, Los Angeles, CA 90095-1606 bprigge @ucla.edu ABSTRACT Prunus eremophila (subgenus Emplectocladus Torr.) is described and illustrated from the southern Mojave Desert. It is closely related to P. havardii (W. Wight) S. C. Mason but differs from it in pubescence of leaves and larger size of floral parts and fruit. Key Words: Prunus, Prunus eremophila, Rosaceae, Amygdaloideae. During vegetation surveys of the Mojave Desert by the Bureau of Land Management (BLM), some unusual specimens of Prunus were collected by BLM botanists. After reviewing specimens of other woolly fruited species of Prunus at LA, RSA, GH, and UC and borrowing specimens of P. havardii from SRSC, it is concluded that they merit recog- nition as a new species. Prunus eremophila Prigge, sp. nov. (Fig. 1).— TYPE: USA. California, San Bernardino Co. 12.8 mi NNW of Goffs on Lanfair Road and 0.8 mi E on road to True Blue Mine, 1145 m elev., 19 March 1992, Prigge 9825 (Holotype: RSA; Isotypes: CAS, GH, LA, MO, NY, TEX, UC). Pruno havardii similis sed differt foliis villosis (non glabris vel subtiliter pubescentibus), hypanthio 3—6 mm longo (vs. 2.0—2.5 mm), sepalis 1.0—1.6 mm longis (vs. 0.5—1.0 mm), petalis 4—7 mm longis (non 2 mm longis vel carentibus), fructu lanato (non pubescenti vel canescenti) 10-16 mm long (vs. 7-10 mm) 9-11 mm lato (vs. 8—9 mm), et en- docarpio 9-14 mm longo (vs. 6—9 mm). Dioecious, + globose to widely spreading, intri- cately branched shrubs to 2.2 m tall; outer bark on older stems gray glaucous becoming reddish brown; inner bark orange; branchlets grayish pu- bescent, weakly thorny but without sharp, indurated tips; internodes 3—14 mm long; short-shoot spurs 1.5—5 mm long. Leaves conduplicate in bud, spath- ulate to ovate, 5-20 mm long and 2—8 mm wide on short shoots, 13—30 mm long and 7-19 mm wide on long shoots, serrate (rarely entire or wavy) with a total of O—13 teeth that are often asymmetrically disposed, sparsely villous on both surfaces, cuneate at base, acute, round, truncate, or retuse, often mu- cronulate at apex, lacking stomata on adaxial sur- face. Petioles 0.5—3.0 mm long on short shoot leaves, 2.0-5.0 mm long on long shoot leaves. Flowers axillary with 2—4 flowers per leaf axil, pre- cocious, unisexual by abortion of either stamens or pistil. Hypanthium turbinate, pubescent externally, strigose internally in male flowers, glabrous inter- nally in female flowers. Sepals 5, 1.1—1.6 mm long, 1.4-2.3 mm wide, deltate, externally pubescent. Petals 5, white, 2.7-5.8 mm long, 2.2—4.0 mm wide, round to spathulate, slightly narrower and more acute in male flowers. Stamens 10—15, in 3 whorls of 5 stamens each or fewer by abortion, the uppermost whorl opposite the calyx lobes and orig- inating on the hypanthium rim, the middle whorl opposite petals, the lowermost whorl + opposite the calyx lobes, with the lower two whorls of sta- mens arising from wall of hypanthium; filaments 1—2.6 mm long, white; anthers 0.7—1.2 mm long, 0.7—1.0 mm wide, light yellow. Pistil one (but oc- casionally two and then connate at the ovary), 4.8— 6.0 mm long, the ovary 1.6—2.5 mm long, 1.5—1.8 mm in diameter, pubescent, the style 3.2—3.7 mm long. Fruit drupaceous, slightly laterally com- pressed ovoid to orbicular, (9—)11—16 mm long, (6—)8—11(—14) mm wide along major axis, (6—)7— 10(—14) mm along minor axis, densely villous with grayish or rusty hairs; sarcocarp (exocarp + me- socarp) very slightly fleshy, ca 0.5 mm thick, easily separating from pit in fertile fruits but generally not split along the ventral suture, exocarp apricot color when ripe. Endocarp smooth, apiculate, and ven- trally ridged, 9-15 mm long, 6—10 mm wide. Paratypes. U.S.A. California. San Bernardino Co.: 12.8 road mi NNW of Goffs along Lanfair Road, then 0.8 mi E on trail-road toward Vontrigger Hills, 1160 m, 19 March 1992, Henrickson 22633 (CSLA); southern Lanfair Valley E of Bobcat Hills, 0.8 mi NE of True Blue Mine, 1145 m, 21 Mar 1980, Prigge 3634 (LA); Vontrigger Hills, 12.8 mi NNW of Goffs on Lanfair Road and ca. 1.95 air mi E of junction of Lanfair Rd and road to True Blue Mine, 1155 m, 3 Apr 1985, Prigge 6235 (LA, RSA); (same area) 7 May 1985, Prigge, Thompson, and Davis 6374 (NY, TEX, UC, RSA, CAS, GH); (same area) 19 March 1992, Prigge 9826 (TEX, RSA); (same area and date), Henrickson 22632 (CSLA); Lanfair Valley, 12.9 air mi N of Goffs 286 MADRONO [Vol. 49 WANs ZZ ! MD WZ Fic. 1. Prunus eremophila: A) fruiting branch with short-shoot leaves (Prigge 6372); B) branch with long-shoot leaves and stipule; C) short-shoot leaves; D) male flower—early anthesis (left), dissected flower with rudimentary pistil (right), and petal (above); E) dissected male flower with two rudimentary pistils (abnormal); F) terminal branch with female flowers and rudimentary stamens; G) dissected female flower with rudimentary stamens; H) endocarp and fruit. where Old Government Road crosses Sacramento Wash, 1165 m, 13 May 1992, Prigge and Henrick- son 10220 (RSA). Prunus eremophila occurs primarily in washes but also on rocky slopes in the higher zones of the creosote bush series and the transition zone from creosote bush to the blackbush series where there are scattered Joshua trees (Sawyer and Keeler-Wolf 1995). Substrates are derived from igneous rocks (granites and rhyolites). Common associates are Te- tradymia_ stenolepis, Ericameria cooperi, Yucca brevifolia, Y. schidigera, Acamptopappus sphaero- 2002] 115° 10' PRIGGE: PRUNUS EREMOPHILA 287 115° 10' LANFAIR VALLEY FENNER VALLEY | to Goffs 10 km Fic. 2. cephalus, Eriogonum fasciculatum, Larrea triden- tata, Salazaria mexicana, Yucca baccata, Lycium oligosperma, Krameria erecta, and Xylorhiza tor- tifolia (nomenclature follows Hickman 1993). Ele- vation ranges from 975-1175 m (3200-3850 ft). The species is known only from the vicinity of the Vontrigger Hills, Bob Cat Hills, southern Lanfair Valley, and one outlier in northeastern Fenner Val- ley of southeastern San Bernardino County (Fig. 2). Flowering time is from mid March to early April. Prunus eremophila is in the subgenus Emplec- tocladus Torr. (commonly included in subgenus Amygdalus L.) as envisioned by Mason (1913). The subgenus is characterized by dioecy through abor- tion of stamens or pistils, a fine pubescence on the inside of the hypanthium, stamen number reduced to 10 or 15 with some filaments inserted on the hypanthial wall, and with or without stomata on the upper leaf surface (Emplectocladus and Prunus are the only subgenera of Prunus that have species without stomata on the upper leaf sufaces). All these traits, including the absence of adaxial leaf stomata, occur in P. eremophila. When Mason studied this subgenus, only four species (all North American) were known: Prunus fasciculata (Torr.) Gray, P. microphylla (H.B.K.) Hemsl., P. minuti- flora Englem., and P. havardii. With the description of P. cercocarpifolia from the Chihuahuan Desert region by Villarreal (1989) and now P. eremophila, the subgenus now consists of six species. A comparison of vegetative, floral, and fruit characters among the species of Emplectocladus point to a possible close relationship between Pru- Known distribution of Prunus eremophila (@) in eastern San Bernardino County, California. nus eremophila and P. havardii of the Chihuahuan Desert of southwest Texas and northern Mexico and, to a lesser degree, with P. microphylla of cen- tral Mexico (states of San Luis Potosi, Hidalgo, México, and District Federal). Prunus havardii is thorny and has glabrous or sparsely pubescent leaves, and P. eremophila is only rigidly branched, not truly thorny and has villous leaves. The larger flower size (longer hypanthium length, longer pet- als, stamens, and pistils) and slightly larger fruit size of P. eremophila are distinctive within the sub- genus Emplectocladus indicating that P. eremophi- la is distinctive and recognizable as a distinct spe- cles. Prunus eremophila is sympatric with P. fasci- culata but flowering time overlaps only partially. The former begins flowering about two to three weeks earlier than the latter, so that P. eremophila has almost completed flowering by the time P. fas- ciculata begins. There is a slight overlap in flow- ering (perhaps as much as a week) when cross pol- lination could occur between these two sympatric species, but no hybrids were found suggesting that additional isolating mechanisms other than seasonal ones are present. At the present, the species does not appear to be threatened, but any species with such a limited dis- tribution and small size (perhaps ca. 2000 individ- uals) could become threatened fairly quickly from off-road vehicle (ORV) use, grazing pressure, and mining activities or global warming. All known populations are within the recently formed East Mojave National Preserve, and the Preserve should 288 minimize or eliminate most threats from human ac- tivities. Future ecological and physiological studies to determine the population size and structure, seed germination requirements, seedling survival, re- cruitment, and threats would provide valuable data for evaluating the long term survivability of this species. Prunus eremophila can be identified by making the following modification to the key in The Jepson Manual (Hickman 1993): 5’ Twigs rigid; ovary and fruit generally densely pu- berulent. 9. Flowers perfect; leaf blades glabrous (some- times minutely puberulent on petiole, midrib, and basal margin in P. fremontii) and more than 3 mm wide. 10. Leaf blades ovate to round, 7—22 mm wide, base obtuse to + cordate; petals WnGer sae. et eye a ee oe P. fremontii 10’ Leaf blades elliptic to oblanceolate, base tapered; petals reddish .... P. andersonii 9’ Flowers imperfect; leaf blades glabrous to vil- lous, if glabrous the leaf blades less than 3 mm wide. 11. Leaf blades spathulate, 1-3 mm wide; puberulent or glabrous .... P. fasciculata 12. Leaf surface puberulent; inland ... var. fasciculata eh Ge) jem ce uleh se) lel se” jel *s¥ fen ce] fe; lelurstiae MADRONO [Vol. 49 12’ Leaf surface glabrous to low-papil- latesycoastall ea ae var. punctata 11’ Leaf blades ovate, 2-19 mm wide, sparsely villous P. eremophila ACKNOWLEDGMENTS I thank Hyrum Johnson for bringing this plant to my attention, Henry J. Thompson, Stan Davis, and James Henrickson for their help and advise on this paper and their company on field trips, Guy Nesom for the Latin discription, Bobbi Angell for the illustration, the curator of the Sul Ross Herbarium for the loan of Prunus havar- dii, the curators of GH and UC/JEPS for permitting me to examine the specimens in their herbarium, and the review- ers (J. Henrickson and Richard Lis) for constructive and helpful comments. LITERATURE CITED HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher plants of California. University California Press, Berkeley, CA. Mason, S. C. 1913. The pubescent-fruited species of Pru- nus of the southwestern states. Journal of Agricultural Research 1:147—177. SAWYER, J. O. AND T. KEELER-WoLFr. 1995. A manual of California vegetation. California Native Plant Socie- ty, Sacramento, CA. VILLARREAL Q., J. A. 1989. A new species of Prunus sub- genus Amygdalus (Rosaceae) from Coahuila, México. Sida 13:273—275. Maprono, Vol. 49, No. 4, pp. 289-294, 2002 A NEW CEANOTAUS (RHAMNACEAE) SPECIES FROM NORTHERN BAJA CALIFORNIA, MEXICO STEVE BOYD Herbarium, Rancho Santa Ana Botanic Garden, 1500 N. College Ave., Claremont, CA 91711 Steve.Boyd@cgu.edu JON E. KEELEY Herbarium, Rancho Santa Ana Botanic Garden, 1500 N. College Ave., Claremont, CA 91711 U.S. Geological Survey, Western Ecological Research Center, Sequoia Field Station, Three Rivers, CA 93271 Department of Organismic Biology, Ecology and Evolution, University of California, Los Angeles, CA 90095 ABSTRACT Ceanothus bolensis S. Boyd & J. Keeley is a new species in the subgenus Cerastes from northwestern Baja California, Mexico. It is well represented at elevations above 1000 m on Cerro Bola, a basaltic peak approximately 35 km south of the U.S./Mexican border. It is characterized by small, obovate to oblan- ceolate, cupped, essentially glabrous leaves with sparsely toothed margins, pale blue flowers, and globose fruits lacking horns. Principal components analysis on morphological traits shows it to be distinct from other members of Cerastes which are distributed away from the coast in southern California and Baja California, Mexico. These phenetic comparisons also suggest that Ceanothus otayensis should not be subsumed under C. crassifolius, as treated in the Jepson Manual, but rather should be retained at specific rank as well. RESUMEN Ceanothus bolensis S. Boyd & J. Keeley es una nueva especie del subgénero Cerastes en el noroeste de Baja California, México. Esta bien representada en altitudes por arriba de los 1000 m en el Cerro Bola, un pico basdltico, apr6ximadamente a 35 km de la frontera de USA/México. Esta se caracteriza por hojas pequenas, obovadas a oblanceoladas, convexas, esencialmente glabras y con margenes espar- cidamente dentados, flores azul palido y frutos globosos sin corniculos. El andlisis de componentes prin- cipales con caracteres morfoldgicos la presenta como distinta de otros miembros de Cerastes distribuidos lejos de la costa en el sur de California y de Baja California, México. La comparaci6on fenética también sugiere que Ceanothus otayensis no debe ser incluida dentro de C. crassifolius como en el Manual de Jepson, sino mas bien deberia ser retenida a nivel especie. Key Words: Ceanothus, subgenus Cerastes, Baja California, Mexico, phenetic analysis, endemic. Ceanothus is a diverse genus of shrubs with the center of species diversity in the Mediterranean-cli- mate California Floristic Province (Schmitt 1993: source for nomenclature, except where indicated). It comprises approximately 60 species (van Rens- salaer and McMinn 1942) more or less equally di- vided into two clades that have long been system- atically recognized as subgenera Ceanothus and Cerastes (Harding et al. 2000). Recent collections of a Ceanothus from north- ern Baja California, Mexico suggest a new taxon that is worthy of recognition. These collections are from Cerro Bola, a basaltic mountain, ap- proximately 35 km south of the Tecate border crossing. At elevations between 1000 m and the peak at 1290 m, a small-leaved, erect, divaricate- ly branched Ceanothus species in the subgenus Cerastes is frequent in chaparral dominated by Arctostaphylos glandulosa Eastw. ssp. adamsii (Munz) Munz and Chamaebatia australis (Bran- degee) Abrams. It is less frequent on lower slopes from at least 500 m elevation. This Ceanothus shares features with several other Cerastes spe- cies distributed in southern California and adja- cent Baja California, Mexico. The lack of fruiting horns and deeply concave, toothed leaves sug- gests an affinity with Ceanothus greggii A. Gray var. perplexans (Trel.) Jepson and with the rela- tively recently described C. ophiochilus Boyd, Ross, & Arnseth (Boyd et al. 1991), a rare en- demic in southwestern Riverside County. The small leaf size of the Cerro Bola plants is similar to C. ophiochilus and C. otayensis McMinn (van Renssalaer and McMinn 1942), a localized en- demic on two mountain peaks in southern San Diego County, California and immediately adja- 290 MADRONO cent Baja California, ca. 30—35 km northeast of Cerro Bola. In the most recent floristic treatment C. otayensis is treated as a hybrid derivative of C. crassifolius Torr., a widespread species away from the coast in southern California and Baja California, and C. greggii var. perplexans, the principle Cerastes species from the interior slopes of the Peninsular Ranges (Schmitt 1993). METHODS Herbarium specimens of the Cerro Bola taxon and other Ceanothus species in subgenus Cerastes from southern California were used for phenetic comparisons. Because fruit morphology has more distinguishing characters than flowers, only fruiting specimens were selected. Species used for compar- ison were Ceanothus greggii var. perplexans, C. greggil var. vestitus (Greene) McMinn, C. ophioch- ilus, C. otayensis, and C. crassifolius. We selected 14 characters for analysis; 6 con- tinuous quantitative, 7 qualitative and 1 calculated ratio (Table 1). For quantitative characters, two samples were measured for each specimen and the mean was used in the analysis. For qualitative characters, characteristics were given a relative score from 1 to 5. All character states were stan- dardized as z-scores by subtracting each observa- tion from the mean of all individuals, and dividing by the standard deviation. A species matrix of these scores was used for ordination with principal components analysis using SYSTAT 5.05 (Evans- ton, IL). RESULTS Means values for phenetic characters for all taxa discussed above are shown in Table 1. The Cerro Bola specimens have the smallest leaves but they are not significantly different from the other two small-leaved taxa, C. ophiochilus and C. otayensis. Cerro Bola plants are similar to C. ophiochilus in their very deeply concave leaves, limited pubes- cence, and smaller fruits that lack horns, however, these two taxa differ in their leaf shape, reflected in the leaf length/width ratio. The broad leaves of Cerro Bola plants are quite unlike the nearly linear leaves of C. ophiochilus, which generates a length/ width ratio roughly double that of all other taxa. The low apical angle on leaves of Cerro Bola plants is quite unlike C. ophiochilus but similar to C. otay- ensis and C. greggii var. perplexans. Ceanothus otayensis separates from all other taxa, except C. crassifolius, in having revolute leaves, well devel- oped, often brownish pubescence on branchlets and undersides of leaves. The principal components analysis for all taxa (Fig. 1), explained 50% of the total variance on the factor 1 axis and 20% on the factor 2 axis. Cea- nothus crassifolius was widely separated from the other taxa on the factor 1 axis; the most important components being convex leaves, revolute margins, LEAF AND FRUIT COMPARISON OF CEANOTHUS BOLENSIS AND OTHER SPECIES IN SUBG. CERASTES (X + SD). TABLE 1. C. greggii var. vestitus C. greggii vat. C. otayensis C. crassifolius C. ophiochilus perplexans C. bolensis Character 10 23.0 + 4.4 WAS 2 2 12 Ply ee eg Sid) ee wile? 12 Bo) 25 OL 26 49+ 0.8 39) ae (0),7/ n 1.9 + 0.3 1.4 + 0.2 32 2 (0,5) 3.8 + 0.8 3,5) ae (04! 522i SG Of aes to 22 OLS) 32) Be il 10.1 + 2.4 Seo) 28) Wnts 1.8 + 0.4 358) SE (08) 10+0 le} Be iol Sia 22 (ils 54.6 + 17.3 14.7 + 2.6 2 22 OJ 2.9 + 0.6 WA, = 9) 1.3 2 O2 4.3 + 0.7 10+ 0 3.4 + 0.7 48 + 5.1 50.1 + 13.6 Leaf length (mm) Leaf width (mm) Length/width ratio 1.0 + 0.0 4.6 + 0.7 Pigs oe Ilsil 2S)S) 22 115).3 SOON 4.5 + 0.5 extreme) extreme) Leaves concave (1 1.0 + 0.0 Io 26 @2 3.8 + 0.6 6.7 + 4.3 ilo) 22 AO no, 5 = none, 5 Leaves revolute (1 = Teeth on leaves (1 53) 22 (0)07/ DY) BET 3} many) Wifes) 2a) Ahn Pubescence on branchlets Apical angle of leaves (°) Basal angle of leaves (°) Boo} 26 (0),2) 3.6 + 0.5 2M 28 (0).3) 44+ 0.5 D0) 22 OO 2.6 + 0.5 dense) glabrous, 5 = él Pubescence on leaf abaxial side [Vol. 49 Le 2e 10) 2s] 0,5 1.0 + 0.0 4.3 + 0.5 49+ 0.3 1.0 + 0.0 glabrous, 5 = dense) ad 2002] BOYD AND KEELEY: Factor 2 -1.5 -1 -.5 0 y 1 Factor 1 CEANOTHUS BOLENSIS 291 C. bolensis C. crassifolius C. greggii perplexans C. g. vestitus C. ophiochilus C. otayensis BOS ae © co 2 Fic. 1. Principal components analysis of Ceanothus crassifolius, C. greggii var. perplexans, C. greggii var. vestitus, C. otayensis, C. ophiochilus, and C. bolensis. Factor loading scores are in Table 2. leaf length, pubescence characters and fruit horns (Table 2). Ceanothus otayensis was clearly sepa- rated from C. crassifolius on both the factor 1 and factor 2 axes but overlapped with C. greggii var. perplexans (Fig. 1). Taking C. crassifolius out of the analysis gave greater resolution to the separation of C. otayensis and C. greggii var. perplexans (Fig. 2). In this analysis, factors 1 and 2 explained 36% and 21%, respectively of the total variance. Ceanothus otay- ensis WaS most prominently separated along the factor 1 axis, where pubescence characters and de- gree of leaf concavity were among the important components. On the factor 1 axis the Cerro Bola plants were distinctly intermediate to C. otayensis and C. ophiochilus. Cerro Bola plants were dis- tinctly separated from C. greggii varieties on the factor 2 axis, where the important components were pedicel length, fruit horns, leaf length and apical angle (Table 2). Clearly there is a sound morphological basis for the recognition of the Cer- TABLE 2. FIGURES 1 AND 2. ro Bola taxon (Figs. 1, 2), treated here as a new species. SPECIES TREATMENT Ceanothus bolensis S. Boyd and J. Keeley, sp. nov. (Fig. 3)—-TYPE: MEXICO, Baja California, chaparral with Chamaebatia australis on NE side of Cerro Bola, S of Tecate, elevation 1000 m, 26 Apr 1996, Jon E. Keeley 27233 (RSA). Differt a C. ophiochilus foliis late obovatis vel late oblanceolatis; a C. otayensis foliis glabris, non concavis, marginibus revolutis; a C. greggii var. perplexans foliis glabris, parvulis (sub 6 mm lon- gis). Erect, divaricately branched shrub, 1—1.5 m tall, lacking basal burl and not resprouting after top- killed. Older stems ashy gray, intricately branched with rigid diverging branches. Younger branches reddish gray and lightly puberulent. Stipules thick- CHARACTERS USED IN PHENETIC ANALYSIS AND FACTOR LOADINGS FOR PRINCIPAL COMPONENTS ANALYSIS FOR Figure | Figure 2 Character Factor 1 Factor 2 Factor 1 Factor 2 Leaf length 0.88 0.04 0.64 0.60 Leaf width 0.78 = | Pee 0.71 0.30 Length/width ratio —0.08 0.86 —0.504 0.460 Leaves revolute 0.83 —0.06 0.66 —0.39 Leaves concave Sos 0.04 —0.78 —0.24 Teeth on leaves 0.11 —0.84 0.52 ae Apical angle of leaves 0.35 OTF ==) Wes, 0.73 Basal angle of leaves =0.22 0.70 —0.44 0.17 Pubescence on branchlets 0.88 —0.17 0.84 —0.10 Pubescence on bottom of leaves 0.88 —0.03 0.77 —0.11 Pubescence includes brown hairs 0.78 tS 0.66 —0.41 Pedicel length 0.67 0.31 0.36 0.81 Fruit width 0.76 0.13 0.42 0.17 Presence of fruit horns 0.82 0.27 0.49 0.63 292 Factor 2 -3 -2.5 -2 -1.5 -1 -5 O .5 Factor 1 RiGs 2: 1. Factor loading scores are in Table 2. ened and persistent, waxy or corky, dark reddish brown to purplish black. Leaves evergreen, thick, coriaceous, yellowish green, opposite, often clus- tered on short axillary spur branches; petioles 0.5— 1 mm long, + 0.4 mm wide, minutely puberulent; blades broadly obovate to oblanceolate, deeply con- cave, (2.5)4—6(10) mm long and (3)4—7(9) mm wide, surfaces sparsely puberulent when young, glabrate in age, margins not revolute, sharply den- tate distally with (1)2—3 pairs of lateral teeth and an apical tooth; midvein prominent abaxially, lat- eral veins obscure, 3—5 pairs. Inflorescence a sub- umbellate axillary raceme, peduncle + 2 mm long, densely short puberulent, bearing 6—8 flowers. Ca- lyx, including receptacular disk + 5 mm wide at anthesis, lobes 5, pale blue, fading cream-white, 1.8—2 mm long, deltoid to ovate, apex acute. Petals 5, pale blue, fading cream-white, 1.8—2 mm long, ladle-shaped, + equally divided into deeply saccate distal blade and filiform proximal claw. Stamens 5, filament 1.8—2 mm long, anther + 0.4 mm. Ovary with style 1.8—2 mm long, 3-lobed, the lobes + 0.4 mm; fruit a globose to depressed globose capsule, dark green to reddish, 3—4 mm diameter, smooth, lacking apical horns, lateral valve crests absent or vestigial. Distribution. At present, Ceanothus bolensis is known only from the mid- to upper slopes of Cer- ro Bola (500 m), where it is locally common in the chaparral vegetation. To our knowledge, no other member of subgenus Cerastes is found on the mountain, and plants are uniform in overall gross morphology. Cerro Bola is noteworthy as a station for several other phytogeographically in- teresting taxa, such as the near-endemic Arcto- staphylos bolensis P. V. Wells, as well as Ceano- thus papillosus Torr. var. roweanus McMinn and Lepechinia cardiophylla Epling, two taxa consid- erably disjunct from their previously know occur- rences in the Santa Ana Mountains of Orange and MADRONO [Vol. 49 C. bolensis C. greggii perplexans C. g. vestitus C. ophiochilus IX ar © C. otayensis Principal components analysis without Ceanothus crassifolius, but including all other taxa considered in Fig. Riverside counties, California (Boyd et al. in prep). Paratypes. MEXICO, Baja California, chaparral with Chamaebatia australis on NE side of Cerro Bola, S of Tecate, elevation 1000 m, 26 Apr 1996, Jon E. Keeley 27232 (BCMEX); ibid, Jon E. Keeley 27227 (CAS); ibid, Jon E. Keeley 27234 (SD); ibid, Jon E. Keeley 27238 (US); ibid, Jon E. Keeley 27236 (MEXU); occasional in chaparral on north slope of Cerro Bola, seen to summit (1275 m), near 31°19.5'N, 116°40’W, elevation ca. 550 m, 6 Jun 1970, Reid Moran 17780 (RSA, SD). RELATIONSHIPS Ceanothus bolensis shows a marked morphologi- cal similarity to several other members of subgenus Cerastes in the southern California region. It is most distinct from the widespread C. greggii var. perplex- ans by having very small leaves that are deeply con- cave. These two leaf traits bear a strong resemblance to C. ophiochilus, however, leaf shape is markedly different, with the latter species being more similar in leaf shape to C. greggii var. vestitus. Ceanothus bolensis is quite distinct from another local endemic, C. otayensis, a taxon restricted to a few mountain peaks about 30-35 km northwest of Cerro Bola. Ceanothus otayensis has been sub- sumed under C. crassifolius (Schmitt 1993) due to the presence of several shared morphological sim- ilarities (revolute leaves and dense pubescence that includes brown hairs). Munz (1959) likewise treat- ed C. otayensis as C. xotayensis McMinn, and sug- gested it was probably a hybrid between C. cras- sifolius and C. greggii var. perplexans. However, principal component analysis on all 14 traits shows C. otayensis to be quite distinct from C. crassifolius (Fig. 1) and distinct from C. greggii (Fig. 2). Ex- tensive exploration of both known southern Cali- fornia localities of C. otayensis, the upper slopes of Otay Mtn. (1090 m) and San Miguel Mtn. (780 m), 2002] BOYD AND KEELEY: CEANOTHUS BOLENSIS 293 Fic. 3. Ceanothus bolensis. A. Fruiting branch showing characteristic small, toothed leaves, and hornless fruits on short axillary peduncles. B. Detail of individual flower. C. Detail of mature capsule showing absence of apical or lateral horns. (Illustrations by Melanie Baer-Keeley.) failed to uncover populations of either C. greggii or C. crassifolius. Ceanothus otayensis forms ho- mogenous populations that appear to be breeding true and lack any indication they are unstable hy- brid swarms (J. Keeley unpublished observations). In addition to the naming of C. bolensis, we suggest C. otayensis be recognized at the specific rank as described by McMinn (van Rensselaer and Mc- Minn 1942). ACKNOWLEDGMENTS Fieldwork and collections by Keeley were conducted in collaboration with Dr. Jose Delgadillo, Universidad Au- tonoma de Baja California, Ensenada, Baja California, Mexico (BCMEX). We wish to extend our thanks to Me- lanie Baer Keeley for preparing the illustration of C. bol- ensis, to Rosa Cerros Tlatilpa for kindly providing the Spanish resumen, and to Elizabeth Friar for assistance in producing figures | and 2. 294 LITERATURE CITED Boypb, S., T. Ross, AND L. ARNSETH. 1991. Ceanothus ophiochilus (Rhamnaceae): a distinctive, narrowly endemic species from Riverside County, California. Phytologia 70:28—41. Haropic, T. M., P. S. SOLTIS, AND D. E. Sottis. 2000. Di- versification of the North American shrub genus Ce- anothus (Rhamnaceae): conflicting phylogenies from MADRONO [Vol. 49 nuclear ribosomal DNA and chloroplast DNA. Amer- ican Journal of Botany 87:108—123. ScHMIpT, C. L. 1993. Ceanothus. Pp. 932-938, 939, 941, 943 in J. C. Hickman (eds.), The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. VAN RENSSALAER, M. AND H. E. McMInn. 1942. Ceano- thus. Santa Barbara Botanic Garden, Santa Barbara, CA. MApRONO, Vol. 49, No. 4, pp. 295-297, 2002 NOTE COLLINSIA ANTONINA IS EVOLUTIONARILY DISTINCT FROM C. PARRYI (SCROPHULARIACEAE SENSU LATO) BRUCE G. BALDWIN Jepson Herbarium and Department of Integrative Biology, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, CA 94720-2465 bbaldwin @uclink4.berkeley.edu W. ScotrT ARMBRUSTER Department of Biology, Norwegian University of Science and Technology, N-7491 Trondheim, Norway and Institute of Arctic Biology, University of Alaska, Fairbanks, AK 99775-7000 A diminutive blue-eyed Mary from shale expo- sures in the San Antonio Hills (Santa Lucia Range) of southern Monterey County, California, was de- scribed by Hardham (1964) as Collinsia antonina. Plants assigned to C. antonina resemble members of C. parryi A. Gray, from the Transverse Ranges of southern California, and have been treated as either close relatives (Munz 1968) or, most recent- ly, as members (Neese 1993) of C. parryi. The con- siderable disjunction between populations in the Santa Lucia and Transverse ranges, although not unique [e.g., Syntrichopappus lemmonii (A. Gray) A. Gray (Compositae)], warrants closer scrutiny. In our evolutionary investigations of Collinsieae, we have examined phylogenetic relationships among taxa throughout the tribe using sequences of the internal transcribed spacer (ITS) region of nu- clear ribosomal DNA (see Armbruster et al. 2002). Upon including plants referable to C. antonina [E. C. Neese 21500 (JEPS)] in our analyses, we were surprised to find that members of C. parryi sensu stricto (s.s.) [E. C. Neese 21530 (JEPS)] appear to be more closely related to members of C. concolor Greene [E. C. Neese 21539A (JEPS)] than to mem- bers of C. antonina. Collinsia concolor and C. par- ryi S.S. constitute a strongly supported (99% boot- strap) clade, to the exclusion of C. antonina (Fig. 1). Collinsia concolor and C. parryi s.s. are both endemic to the southern California Floristic Prov- ince and differ greatly in inflorescence architecture and flower size (Table 1). Based on the tree topol- ogy presented in Fig. 1, small flower size may be a shared, ancestral characteristic of C. antonina and C. parryi, with large flowers being a derived feature of C. concolor. Such major shifts in flower size have occurred repeatedly throughout the evolution- ary history of Collinsia (Armbruster et al. 2002). Evolutionary distinctiveness of C. antonina and C. parryi s.s. 1s also reflected by differences in morphological characteristics (Table 1). In keys to species of Collinsia (e.g., Newsom 1929), C. an- tonina might be confused with the distantly related C. childii Parry ex A. Gray because both taxa have glandular inflorescences and small flowers and both occur in the Santa Lucia Range. The two taxa can be readily distinguished by differences in seed number per capsule (Table 1). Hardham (1964) recognized two sympatric subspecies of C. anton- ina (C. a. subsp. antonina and C. a. subsp. pur- purea Hardham), which await further systematic evaluation. Collinsia antonina warrants recognition as a dis- tinct species based on the above molecular and morphological evidence. This narrowly endemic taxon was earlier considered rare and endangered by the California Native Plant Society (CNPS) (Smith and Berg 1988); since 1993, CNPS has fol- lowed Neese’s (1993) treatment of Californian members of Collinsia by including C. antonina within the circumscription of the common species, C. parryi (Skinner and Pavlik 1994; California Na- tive Plant Society 2001). In light of our findings, C. antonina deserves renewed attention by plant conservationists. ACKNOWLEDGMENTS We thank Elizabeth Chase Neese for her invaluable field collecting, taxonomic insights, and review of the manuscript, Bridget L. Wessa for extensive laboratory as- sistance, and reviewers Noel H. Holmgren, David J. Keil, and John L. Strother for helpful suggestions. This research was supported by grants from the Lawrence R. Heckard Endowment Fund of the Jepson Herbarium (to BGB), the Norwegian Research Council (to WSA), and NSF DEB- 9708333 (to WSA). 296 MADRONO 100 dg 71 50 d2 59 d3 97 d6 100 ds 97 d7 76 d3 65 a3 100 d1i3 65 d2 Tonella tenella T. floribunda [Vol. 49 Collinsia heterophylla C. greenei C. sparsiflora C. bartsiifolia C. corymbosa C. tinctoria C. parryi C. concolor C. antonina C. multicolor C. callosa C. childii C. parviflora C. grandiflora C. verna C. violacea C. rattanii C. linearis lA C. torreyi var. torreyi C. torreyi var. wrightii Fic. 1. The most parsimonious tree from phylogenetic analysis of nuclear ribosomal DNA sequences of the internal transcribed spacer region in tribe Collinsieae (Baldwin et al. unpublished; see Armbruster et al. 2002). The tree was rooted with sequences from outgroup taxa in tribe Cheloneae (Chelone, Keckiella, and Penstemon). Numbers above branches are bootstrap values (only values = 50% are shown); numbers below branches, preceded by “‘d’’, are decay values (only values > d1 are shown). The clade including Collinsia antonina, C. concolor, and C. parryi is highlighted. TABLE |. SOME MORPHOLOGICAL DIFFERENCES BETWEEN COLLINSIA ANTONINA AND SIMILAR OR CLOSELY RELATED SPECIES. C. antonina C. parryi Inflorescence glandularity glandular eglandular Longest-leaf length 5-10 mm (5—)10—45 mm Pedicel length <10 mm (5—)10—45 mm Corolla length 6—7 mm (5—)7—10 mm Corolla color white or purple lavender-blue Seeds/fruit 6-8 8—12 C. concolor glandular or eglandular 10—45 mm 1-5 mm 10-16 mm blue/lavender— blue and white 11-12 C. childii glandular 10—50 mm 5—25 mm 6-8 mm pale lavender or white 2 2002] LITERATURE CITED ARMBRUSTER, W. S., C. P. H. MULDER, B. G. BALDwWIn, S. KALISZ, B. WESSA, AND H. Nute. 2002. Comparative analysis of late floral development and mating-system evolution in tribe Collinsieae (Scrophulariaceae s. 1.). American Journal of Botany 89:37—49. CALIFORNIA NATIVE PLANT SoctETy. 2001. Inventory of rare and endangered vascular plants of California, 6th ed. Rare Plant Scientific Advisory Committee, D. Ti- bor, Convening Editor. California Native Plant Soci- ety, Sacramento, CA. HARDHAM, C. B. 1964. A new Collinsia from Monterey County, California. Leaflets of Western Botany 10: 133-135. BALDWIN AND ARMBRUSTER: COLLINSIA ANTONINA 297 Munz, P. A. 1968. Supplement to a California flora. Uni- versity of California Press, Berkeley. NEESE, E. C. 1993. Collinsia. Pp. 1024—1027 in J. C. Hickman (ed.), The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. Newsom, V. M. 1929. A revision of the genus Collinsia. Botanical Gazette 87: 260-301. SKINNER, M. K., JR. AND B. M. PAVLIK (eds.). 1994. In- ventory of rare and endangered vascular plants of California, 5th ed. California Native Plant Society, Sacramento, CA. SmiTH, J. P., JR. AND K. BERG (eds.). 1988. Inventory of rare and endangered vascular plants of California, 4th ed. California Native Plant Society, Sacramento, CA. MADRONO, Vol. 49, No. 4, p. 298, 2002 REVIEW Field guide to liverwort genera of Pacific North America, by W.B. Schofield. 2002. Global Forest So- ciety in association with the University of Washington Press, San Francisco, CA. 232 pp., 93 line drawings, glossary, and index. 7” X 10”. $25.00. ISBN 0-295- 98194-6. Available at www.washington.edu/press Most of the liverwort field guides and other tax- onomic treatments covering this group of organ- isms in North America have focused primarily east of the 100th Meridian. It is therefore most welcome that this field guide covers much of the geograph- ical area of interest to Madrofo readers. This pa- perback book, while not quite a size to readily carry in the field, 1s nonetheless not heavy or excessively bulky. What we do have here is a well-designed volume. Bryophytes, the first land plants, are ar- ranged into three lineages. The liverworts (and hornworts) are clearly less well known to most vas- cular-trained botanists than are the mosses. This volume is easy to use and will expand one’s knowl- edge about this group of land plants. In some ways, this is a companion work to Schofield’s field guide of Some common mosses of British Columbia pub- lished back in 1969. The Introduction to the liver- wort genera field guide covers the first 22 pages and is concise yet very informative. The Introduc- tion addresses how to collect liverworts and horn- worts, followed by a short overview of collecting history of liverworts in Alaska, British Columbia, Washington, Oregon and California. Habitats, sea- sonality of finding liverworts, and distribution pat- terns in the region follow. Implications of liver- worts to people end the Introduction section. In most field guides, it is the illustrations and keys that determine how useful the book will ac- tually be, especially to someone approaching the group for the first time. In this regard Schofield’s book shines. While all keys have caveats about their use, I personally found the keys to be rather easy to use. The keys keep jargon and bryological terms to a basic minimum, and the couplets are succinct. The couplet choice is designed without a lot of overlapping characters. In many cases, genera key out in multiple places to accommodate the di- versity of species within larger liverwort genera. The key itself is divided into eight sub-key sections to further expedite the identification process. Each genus is arranged alphabetically so it is easy to lo- cate, and each genus has a full-page illustration. The illustrations are visually attractive and de- signed to represent a genus without being too tech- nical. In some ways, they are more “‘artistic”’ than a detailed illustration, such as would be used in describing a new taxon. Actually, I think the illus- trations provided are more effective as a represen- tation at the genus level. Since liverworts are gen- erally small plants, the bar scale provided with each illustration is important to review so one has a bet- ter idea of the organism’s actual size. A template is provided for each genus so com- parisons can be made readily among closely related genera. Each genus has a short explanation about the meaning of its name, a statement of the number of species in the genus, its habit, its habitat, repro- duction, local distribution, world distribution, dis- tinguishing characteristics and similar genera. The genera of hornworts within the Pacific North America are also included, adding further to the value of this book since one could confuse a horn- wort for a thallose liverwort. However, hornworts upon closer examination can readily be distin- guished from the liverworts, especially when spo- rophytes are present. The determination of the two hornwort genera covered in the book, cccurs at couplet 28 of key Hl. Another interesting feature, and I think a great decision, is adding Takakia (Takakiaceae) to the book. The placement of this genus within the bryophytes has been a consider- able puzzle, with some specialists insisting it is a liverwort while others opted to say it was more closely aligned with the mosses. Although it has been finally determined to be basal to the moss lin- eage (based on the sporophyte), it clearly looks more like a liverwort when only the gametophyte is present. I hope that additional occurrences of this relatively rare genus can be located in the Pacific Northwest. Having the genus illustrated here should be a good first step for others to seek it out. I strongly recommend this new field guide to all botanists with an interest in learning more about our non-flowering plants. As Schofield’s book states, liverworts are a remarkable group of plants, and upon closer observation, they make a consid- erable contribution to forest ecosystems, especially in the temperate rain forests along the coast of western North America. At $25.00 this guide is a good investment. Will you be able to key every liverwort to species with this field guide? Abso- lutely not. But, having the genus properly identified will speed up considerably the time to do so in more detailed bryofloras. This field guide is a won- derful introduction to recognizing genera of liver- worts in Pacific North America. In short order you will be able to recognize the common liverwort genera on rotten wood and logs, soil and humus, tree trunks, rock outcrops and even those liverworts submerged in springs and streams. I recommend it to botanists, ecologists, foresters, and conservation biologists without any reservations. —JAMES R. SHEVOCK, Research Associate, Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. MADRONO, Vol. 49, No. 4, p. 299, 2002 PRESIDENT’S REPORT FOR VOLUME 49 I am pleased to report that 2002 was a year of accom- plishment for the California Botanical Society, thanks to the dedicated efforts of the Editors and Council. Editor John Callaway skillfully concluded his first year in charge of Madrono, with four excellent issues completed. The quality, breadth, and depth of articles published in Ma- drono, Volume 49, testifies to the vibrancy of our journal and the effectiveness of John’s editorship. Thank you, John! I also thank our Book Editor Jon Keeley, Notewor- thy Collection Editors Dieter Wilken and Margriet Weth- erwax, and the Board of Editors for their important con- tributions to Volume 49. Visibility of articles in Madrono was enhanced consid- erably in 2002 by internet access to titles and abstracts for new issues of the journal through the California Botanical Society web-site (www.calbotsoc.org). John Callaway and Society web-master Curtis Clark worked with Allen Press to provide direct links to contents of Madrono (beginning with Vol. 49, issue 1) via Allen Press’s APT Online. The results of their efforts will help to increase citation and impact of articles in Madrono across the international bo- tanical community. Over the last year, web-masters Curtis Clark and John LaDuke have made other major improvements to the So- ciety web-site (www.calbotsoc.org), which has become our main clearinghouse for information about publishing in Madrono and about society activities, events, and mem- bership. Curtis and John developed a new, streamlined organization for the site, which is easily negotiated with well-placed links. The site also contains links to other so- cieties, institutions, and resources of importance to west- ern North American botany. Many thanks, Curtis and John! Thanks to efforts by Corresponding Secretary Sue Bainbridge and Treasurer Roy Buck, the Society can now accept credit-card payment for membership dues, using either the web-site membership forms (for new and re- newing members) or renewal envelopes sent with the third issue of Madronio each year. We trust that the convenience of using a credit card will encourage timely renewal of memberships, which remains an urgent need for the So- ciety. Thanks very much to those members who responded quickly to their renewal notices. For those who have not yet responded, please send your renewals in time for re- ceipt on 28 February 2003. Timely renewal of member- ship saves the Society considerable expenditure of effort (although late renewal is far better than a lapse in mem- bership). Significant growth of our membership base would allow the Society to do more to promote botanical research and education. Toward that end, please continue to encourage your colleagues to join us and to publish in Madrono. Also, you can help to support botanical research in eco- nomically depressed, developing countries by giving a sponsoring membership or subscription to a foreign sci- entist or scientific institution or by donating unused copies of Madrono. For more information on making such a gift, please contact Corresponding Secretary Sue Bainbridge (suebain @sscl.berkeley.edu). Fall 2002 saw outstanding lectures at our monthly meetings in Berkeley by Martin Bidartondo, Truman Young, and Randy Jackson, and we look forward to pre- sentations by distinguished scientists David Ackerly, Kim Steiner, Todd Dawson, and Marcel Rejmanek in winter and spring of 2003. I am indebted to outgoing First Vice- President Rod Myatt for his successful efforts, once again, at assembling such a fine lecture series for the Society’s program year. Attendance at our monthly meetings has been strong again this year and I encourage those of you who have an opportunity to participate to please join us, bring a friend, and inform your colleagues about upcom- ing lectures. I also strongly encourage our members to attend the Society’s biennial graduate student meeting and annual banquet at the University of San Diego on Saturday, 15 February 2003. At the graduate student meeting, held dur- ing the day, students from different institutions will be presenting their research proposals, research-in-progress, or completed research in botany in a standard scientific- meeting format. We look forward to a stimulating day of research ideas and results from the promising, next gen- eration of botanists. In the evening, at the annual banquet, we will have the great pleasure of hearing from renowned Baja floristician and cactus expert Dr. Jon Rebman, who will be presenting an after-dinner lecture on recent floristic discoveries in Baja California, Mexico. Jon’s expeditions to remote, beautiful, and under-explored regions of Baja California, in collaboration with Mexican scientists, are fine examples of American/Mexican cooperation and ac- complishment in science. Thanks very much to Second Vice-President Michael Mayer for planning and organiz- ing the upcoming banquet and to Graduate Student Rep- resentatives Robert Lauri and Elizabeth Zacharias for co- ordinating and conducting the graduate student meeting. Last, but by no means least, I thank Recording Secre- tary Staci Markos and Council Members Jim Shevock, Dean Kelch, and Anne Bradley for their dedicated com- mitment and contributions to furthering the goals of the Society, and to all of our members for your continuing support and participation in the Society’s activities and events and for choosing to submit your botanical manu- scripts to Madrono. This is my last year as President of the Society and I especially thank all of you for making this experience so enjoyable and rewarding for me. Have a great year in 2003! —BRUCE G. BALDWIN December 2002 MADRONO, Vol. 49, No. 4, p. 300, 2002 EDITOR’S REPORT FOR VOLUME 49 This report serves to inform the members of the Cali- fornia Botanical Society of the status of Madrono, from the number of manuscripts submitted to papers published. Since the previous editor’s report (see Madrono 49[4}), the journal has received 61 manuscripts for review, in- cluding Articles, Notes, and Noteworthy Collections; 31 of these manuscripts have been accepted for publication in that same time period. The average time for article sub- mission to publication remains at approximately six months. Accepted manuscripts are typically published within approximately three to four months. Few manu- scripts were rejected after review; authors of Madrono ar- ticles did a fine job of responding to reviewers’ sugges- tions. Over the past five years, there has been a substantial effort to get Madrono back on its regular publication schedule, and we are now very close to being on schedule, with six issues published in 2002 (three from volume 48 and three from volume 49). Kristina Schierenbeck (editor of volumes 45—48) deserves most of the credit for this, as she put in an enormous effort to get the journal back on schedule. In addition to catching up on our publication schedule, other improvements have been made with the journal. As noted by Bruce Baldwin in his President’s Re- port, abstracts of Madrono are now available on-line via the California Botanical Society’s web-site (www. calbotsoc.org). In addition the web-site will soon have more detailed Instructions for Authors for Madrono man- uscripts. With volume 49 we initiated a new policy, en- couraging authors to submit names of two to four potential reviewers for manuscripts. If readers have suggestions for other improvements for the journal please let me know. I want to thank the many people who make Madrono possible and who have been incredibly helpful in editing the journal: Kristina Schierenbeck, who gave me lots of help in taking over the editorship and has always been there to answer my questions about the journal; Bruce Baldwin and other members of the CBS Executive Coun- cil, who are always extremely helpful and supportive of everything related to Madrono; Dieter Wilken and Mar- griet Wetherwax, who handle all of the reviews for Note- worthy Collections (and without whom the Noteworthy Collections would not be possible); Jon Keeley, who does all of the book reviews; Steve Timbrook, who provides the annual index of Madrofo articles and the annual table of contents; the Board of Editors, who have provided in- put and advice whenever I’ve asked (especially Norm Ellstrand and Carla D’ Antonio, who are finishing their terms this year); Annielaurie Seifert at Allen Press, who has been extremely helpful when any editorial question arises; the College of Arts and Sciences at the University of San Francisco, who support my efforts on Madrono; and most of all the Madrono authors, who continue to submit outstanding manuscripts, and the reviewers (see the accompanying list of reviewers), who put in a sub- stantial effort to improve the quality of manuscripts that Madrono publishes. MADRONO, Vol. 49, No. 4, pp. 301, 2002 Frank Almeda George Argus Jayne Belnap Jere Boudell Thomas Boyle Steven Broich Matt Brooks Mark Brunell Steve Caicco Robert Callihan Tina Carlsen Raymund Chan Anita Cholewa Ranessa Cooper Daniel Crawford Curtis Daehler Carla D’ Antonio Gerrit Davidse Chuck Davis Frank Davis Jose Delgadillo Rebecca Dolan Andrew Dyer Erin Espeland Ray Evert Phyllis Faber Richard Felger Wayne Ferren Peggy Fiedler Jack Fisher Ted Fleming Peter Goldblatt Leslie Gottlieb Jason Hamilton Héctor Hernandez Macias Noel Holmgren Kent Holsinger Bryan Jennings Steven Jessup Michael Josselyn Jon Keeley David Keil Seung-Chul Kim Robert Leidy Celi Tim Lowrey Carol Mallory-Smith Maria Mandujano Joe McAuliffe Kimberlie McCue REVIEWERS OF MADRONO MANUSCRIPTS 2002 Dale McNeal Kyle Merriam Norton Miller Timothy Miller Richard Minnich Margaret Moore Lorraine Parsons Bob Patterson Eric Peterson Jeanne Ponzetti Jon Rebman John Reeder Marcel Rejmanek Rhonda Riggins James Shevock Stanley Smith Humberto Suzan Anthony Swinehart Teresa Terrazas Dale Vitt Dieter Wilken Paul Wilson Diana Wolf Lidia Yoshida Peter Zika MApRONO, Vol. 49, No. 4, pp. 302-304, 2002 INDEX TO VOLUME 49 Classified entries: major subjects, key words, and results; botanical names (new names are in boldface); geographical areas; reviews, commentaries. Incidental references to taxa (including most lists and tables) are not indexed separately. Species appearing in Noteworthy Collections are indexed under name, family, and state or country. Authors and titles are listed alphabetically by author in the Table of Contents to the volume. Aceana novae-zelandiae, noteworthy collection from OR, 194. Agavaceae: Agave palmeri, effects of fire on reproductive biology, 1; nomenclatural history of Hesperoyucca whipplei and Yucca whipplei, 20. Agave palmeri, effects of fire on reproductive biology, 1. Agrostis Howellii, noteworthy collection from OR, 194. Alnus rubra, noteworthy collection from MT, 55. Ambrosia: A. includes Hymenoclea, 143. New combinations: A. monogyra, A. Mauseth; Roberto: Kiesins and! Carlosy © sto eizay ee eee Hamilton, Jason G., James R. Griffin and Mark R. Stromberg, Long-term population dynamics of native Nassella (Poaceae) bunchgrasses in central California Haubensak, Karen A. (see Parker, Ingrid M.) Heidel, Bonnie (see Spribille, Toby, et al.) 77) 135 299 eS) 165 201 54 289 130 158 186 54 58 33 198 2002] TABLE OF CONTENTS Hrusa, Fred, Barbara Ertter, Andrew Sanders, Gordon Leppig and Ellen Dean, Catalogue of non-native vascular plants occurring spontaneously in California beyond those addressed in The Jepson Manual—Part I Jessup, Steven L., Reticulate ancestry in Mexican Gaudichaudia (Malpighiaceae) ______.-_-----_------ Jessup, Steven L., Six new species and taxonomic revisions in Mexican Gaudichaudia (Malpighiaceae) Johnson, Phillip (see Rice, Barry A., and Phillip Johnson) Jones, James S. (see Garrison, Barrett A., et al.) Kapiane Divhaldsice. Micticalomoray Ollie tO” oe ee Ee ee Karlik, John F, et al., Eugene D. Albertson, Y. Jae Chung, Alistair H. McKay and Arthur M. Winer, Field assessment of the California GAP analysis program GIS database in central California Keeley, Jon E. (see Boyd, Steve, and Jon E. Keeley) Koehler, Catherine E. (see Montalvo, Arlee M.) Leon de la Luz, José Luis (see Rebman, Jon) Leppig, Gordon (see Hrusa, Fred, et al.) Levine, Larry, Mary Bacca and K. O. Fulgham, Plant zonation in a Shasta County salt spring supporting the nly ows populatouvor Puccmellia howellit (POaCcac) a Makines sblizabeth. Notewormuy COUechHon trom ATIZONA, = 2 Marsh, Glenda D., and Debra R. Ayres, Genetic structure of Senecio layneae (Compositae): A rare plant of the SUELO RINT I Ss es ce OE a = 58 Ne 0) Se eR ek ee he ee ee Mathiasen, Robert L., and Carolyn M. Daugherty, Adult sex ratio of Arceuthobium gillii (Viscaceae) McClaran, Mitchel P., and James W. Bartolome, Noteworthy collections from Moorea, Society Islands, French POLSTREST EL a2 sees eee a ad oa a ee NS eee nO ee McKay, Alistair H. (see Karlik, John F, et al.) Mishler, Brent D. (see Stark, Lloyd R.) Montalvo, Arlee M., Laura J. Feist-Alvey and Catherine E. Koehler, The effect of fire and cold treatments on seed germination of annual and perennial populations of Eschscholzia californica (Papaveraceae) in south- BOE Ge OES oe ee a a a Ee ee pe aera oa ta ie ae Newhouse, Bruce, Richard Brainerd and Peter EK Zika, Noteworthy collections from Oregon _ Nobel, Park S., Erick De la Barrera, David W. Beilman, Jennifer H. Doherty and Brian R. Zutta, Temperature lintraviogs tor Cultivation, of edible cachiim: Califomia«.. 2 ae Se ee Oberbauer, Thomas A. (see Rebman, Jon) Parker, Ingrid M., Alexandra Engel, Karen A. Haubensak and Karen Goodell, Pollination of Cytisus scoparius (Fabaceae) and Genista monspessulana (Fabaceae), two invasive shrubs in California Parker, V. Thomas (see Vasey, Michael C.) Patten, Ann M., and Steven J. Brunsfeld, Evidence of a novel lineage within the Ponderosae Pivorunas, David, Veva Stansell and Peter E Zika, Noteworthy collection from Oregon Prigge, Barry A., A new species of Prunus (Rosaceae) from the Mojave Desert of California Rebman, Jon, Thomas A. Oberbauer and José Luis Leén de la Luz, The flora of Toro Islet and notes on Gradaimpodsiand. Baja Calera Meee er ne ee Paice inv ee INOLewortiny COMCciOnS 1oml) Calitonita,. 0... 0 ete ee ee Oe Rice, Barry A., and Phillip Johnson, Noteworthy collection from Alaska Roché, Cindy Talbott (see Gamarra, Roberto) Sanders, Andrew (see Hrusa, Fred, et al.) Sawyer, John O., Review of J/lustrated Field Guide to Selected Rare Plants of Northern California edited by Gary Nakamura and Julie Kiersteand Nelson Sawyer, John O. (see also Eckert, Andrew J.) SAVE Ie atMCeH NOL WOE COM ec itl: REmiy, VV AS CINTA LOIN a Sp ei Shaw, David C., and Steven A. Acker, Canopy macrolichens from four forest stands in the southern Sierra mixed SOARES CIEE SES Oe PILED cd) CEO SCO EN UNIT AS AEC a ALR 2 cect etter ser once Shevock, James R., Review of Field Guide to Liverwort Genera of Pacific Northwest America by W. B. Schofield Slauson, Liz A., Effects of fire on the reproductive biology of Agave palmeri (Agavaceae) SOLO keaEIN Ole WOOLEY CONCCHONS TtOnk Isao, nes. tr ee ee ce Sproulc, Lou, ct al. Noleworuny collections trom Montana 2 Stansell, Veva (see Pivorunas, David) Stark, Lloyd R., Alan T. Whittemore and Brent D. Mishler, Noteworthy bryophyte records from the Mojave DSSERS oe po Reo a 2S es rr! 2 ee Stromberg, Mark R. (see Hamilton, Jason G.) Strother, John L. (see Greenhouse, Jeffrey A.) Strother, John L., and Bruce G. Baldwin, Hymenocleas are ambrosias (Compositae) Triepke, FE Jack (see Spribille, Toby, et al.) Triggs, Matthew L. (see Garrison, Barrett A., et al.) Tweed, William, Review of Seeing Things Whole: The Essential John Wesley Powell ed. William deBuys ___.. Vanderhorst, Jim (see Spribille, Toby, et al.) Vasey, Michael C., and V. Thomas Parker, Review of The Manzanitas of California, also of Mexico and the Oe PES EU Saas USE OS Dt ied BD ae ie D8 Pe ae 0 see ce Wachs, Robin L. (see Garrison, Barrett A., et al.) Wilson, Barbara L., and Sami Gray, Resurrection of a century-old species distinction in Calamagrostis —_... Winer, Arthur M. (see Karlik, John F, et al.) ill 178 132 150 12 132 207 194 228 189 285 145 193 Whittemore, Alan T. (see Stark, Lloyd R.) York, Dana A., Eriogonum ovalifolium var. monarchense (Polygonaceae), a new variety from the southern Sierra Nevada, California Zika, Peter E, Noteworthy collections from Oregon and Washington Zika, Peter E (see also Newhouse, Bruce) Zika, Peter E (see also Pivorunas, David) Zutta, Brian R. (see Nobel, Park S., et al.) 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