VOLUME 57, NUMBER 4 EAR ¢ & i 8 em pay mom UE Rey se Ry A ee gist 0 FE PS PAPAL oe eet AD enon gape see tenis anette” DENSITY AND ELEVATIONAL DISTRIBUTION OF THE SAN FRANCISCO PEAKS RAGWORT, PACKERA FRANCISCANA (ASTERACEAE), A THREATENED SINGLE-MOUNTAIN ENDEMIC JAMES F. FOWLER AND CAROLYN HULL SIEG Cece eer ercreccsccscsreseeeseseersesreseseseoesoeeecoee PLANT POPULATION AND HABITAT CHARACTERISTICS OF THE ENDEMIC SONORAN DESERT CACTUS PENIOCEREUS STRIATUS IN ORGAN PIPE Cactus NATIONAL MONUMENT, ARIZONA Greta Anderson, Sue Rutman, and Seth M. Munson .......c..ccecccseecceneecenees STAND DEVELOPMENT ON A 127-YR CHRONOSEQUENCE OF NATURALLY REGENERATING SEQUOIA SEMPERVIRENS (TAXODIACEAE) FORESTS Will Russell and Kristin Hageseth Michels ........cccccccceeeeesccscccsseceeeeeeseeees REDISCOVERY OF PLAGIOBOTHRYS HYSTRICULUS (BORAGINACEAE) AND NOTES ON ITS HABITAT AND ASSOCIATES Robert E. Preston, Brad D. Schafer, and Margaret Widdowson .............. TAXONOMIC NOVELTIES FROM WESTERN NORTH AMERICA IN MENTZELIA SECTION BARTONIA (LOASACEAE)) John J. Schenk and Larry Hufford Ceo error ere reer serereeeesereeeereseeesesseeesesereresoeseeoes BRODIAEA MATSONII (ASPARAGACEAE: BORDIAEOIDEAE) A NEW SPECIES FROM SHASTA COUNTY, CALIFORNIA RODE Es LTeStON nace HOWELLANTHUS DALESIANUS, RECOGINITION OF A NEW GENUS AND SPECIES IN TRIBE PHACELIEAE (BORAGINACEAE) Genevieve K. Walden and Robert Patterson ...ccccccccccccccccsscccscccusccnscsusssueess CALIFORNIA OREGON Peer er erreresecerseseseseesoeersesereseserseee Peer ee reereeecreseesesesecseesesesesesesesesseceeseses Cece sec er rere seesccsesoeseseesoesreseseesseeeseeseeeeseeeoee Cece errr sere sersessesoesresesoeseeseseesesesereseseeseseoe® OCTOBER-—DECEMBER 2010 MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. 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Staci Markos, University and Jepson Herbaria, University of California, Berkeley, CA 94720, smarkos @berkeley.edu. Graduate Student Representatives: Ben Carter, Department of Integrative Biology and University Herbarium, University of California, Berkeley, CA 94720, bcarter@berkeley.edu. Webmaster: Susan Bainbridge, Jepson Herbarium, University of California, Berkeley, CA 94720-2465, sjbainbridge @ berkeley.edu. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). MADRONO, Vol. 57, No. 4, pp. 213-219, 2010 DENSITY AND ELEVATIONAL DISTRIBUTION OF THE SAN FRANCISCO PEAKS RAGWORT, PACKERA FRANCISCANA (ASTERACEAE), A THREATENED SINGLE-MOUNTAIN ENDEMIC JAMES F. FOWLER AND CAROLYN HULL SIEG USFS Rocky Mountain Research Station, 2500 S Pine Knoll Drive, Flagstaff, AZ 86001 jffowler@fs.fed.us ABSTRACT Packera franciscana (Greene) W. A. Weber and A. Love is endemic to treeline and alpine habitats of the San Francisco Peaks, Arizona, USA and was listed as a threatened species under the Endangered Species Act in 1983. Species abundance data are limited in scope, yet are critical for recovery of the species, especially in light of predictions of its future extinction due to climate change. This study defined baseline population densities along two transects which will allow the detection of future population trends. Packera franciscana ranged from 3529 to 3722 m elevation along the outer slope transect and densities were 4.18 and 2.74 ramets m ’ in 2008 and 2009, respectively. The overall P. franciscana 2009 density estimate for both transects was 4.36 ramets m * within its elevational range of occurrence, 3471—3722 m. The inner basin density was higher, 5.62 ramets m °, than the estimate for outer slopes, 2.89 ramets m °. The elevation of the 2009 population centroid for both transects was at 3586 (+10 SE) m with the inner basin centroid significantly lower than the outer slopes centroid: 3547 (+7 SE) m vs. 3638 (+7 SE) m, respectively. In mid-September, 6—9% of the P. franciscana ramets were flowering and/or fruiting in 2008-2009. These results and our estimate of >36,000 P. franciscana ramets in <2% of its range would suggest that the species is locally abundant, persisting and reproducing. Key Words: Alpine, narrow endemic, Senecio franciscanus, talus, trail transect. Packera franciscana (Greene) W.A. Weber and A. Love (Asteraceae), San Francisco Peaks ragwort, is endemic to treeline and alpine habitats of the San Francisco Peaks in Arizona (Green- man 1917; Barkley 1968; Trock 2006) (Fig. 1) where it has been reported to mostly occur between 3525 and 3605 m elevation (Dexter 2007) or, more generally, 3200-3800 m (Trock 2006). Packera franciscana predominately inhab- its loose talus slopes (USFWS 1983) and has a reported range size of 85 ha (Dexter 2007). Since the elevation of the highest peak on the mountain is 3854 m, there is little habitat available for the plant to migrate upward in a warming climate scenario, and it has been widely speculated that the species is vulnerable to extinction due to climate change. In 1985, the distribution of P. franciscana on the San Francisco Peaks was mapped (Dexter 2007), but little published data 1s available on species abundance. Packera francis- cana, as Senecio franciscanus Greene (Greene 1889), was listed as a threatened species under the Endangered Species Act by the U.S. Fish and Wildlife Service (1983) due to its narrow geo- graphic distribution, apparent habitat specificity, and concerns over recreational impacts. Four other Packera taxa primarily grow in alpine- subalpine talus habitats in western America and two of those, Packera musiniensis (S. L. Welsh) Trock and Packera malmstenii (S. F. Blake ex Tidestrom) Kartesz, are also of conservation concern (FNA 1993+). The evolutionary rela- tionships of P. franciscana within Packera are unknown. Global mean temperature is predicted to rise 1-6°C in the next century due to increased concentration of greenhouse gases (Solomon et al. 2007) and has increased 0.4°C over the previous 150 yr (Trenberth et al. 2007). The predicted general response of species to this increased warmth is to migrate north in latitude or up in altitude (Grabherr et al. 1994; Theurillat and Guisan 2001; Walther 2002; Root et al. 2003; Kullman 2008; Lenoir et al. 2008). Alpine species population decline and extinction are also possi- ble since there may be little available habitat for species’ upward migration (Chapin and Korner 1994; Grabherr et al. 1994; Theurillat and Guisan 2001). These potential ecological changes indicate the need to establish baseline plant species’ distributions and abundances at local scales to definitively detect changes (Post et al. 2009). Locally abundant, single mountain endemic species offer good opportunities to establish baseline studies for this purpose. Kruckeberg and Rabinowitz (1985) note that narrow endemics can be locally abundant in specific habitats but geographically restricted. Biologists have long observed that P. franciscana is common to abundant in the alpine of the San Francisco Peaks (Greene 1889; Little 1941; Phillips and Peterson 1980; USFWS 1983; Phillips and Phillips 1987; Trock 2006), but peer-reviewed studies of P. franciscana abun- 214 MADRONO [Vol. 57 Humphreys Peak A A Agassiz Peak US Route 180 Fic. |. A Fremont Peak Fl agstaff San Francisco Peaks Study Area N 6 Kilorneters ef een 0 15 3 Bee eet Oe ee eee eee Packera franciscana study area showing the San Francisco Peaks in northern Arizona with the inner basin in an extended volcanic caldera to the northeast. Humphreys Peak is at 35°20'45.781”N; 111°40'40.102”W; and 3854 m elevation. dance do not exist. Yet, such data are critical for recovery of the species under the Endangered Species Act from either recreational impacts or future climate change. Off trail use of above treeline habitats on the San Francisco Peaks is currently banned due to concerns about potential negative impacts on P. franciscana. Our study defines baseline population densities along per- manent transects under current climate and recreational use conditions which therefore may allow the detection of future population trends. Specifically, our objectives are to: 1) establish a statistically robust sampling protocol for long- term population density trends; 2) determine the elevation of patch centroids along these transects to allow early detection of climate change driven altitudinal migration; and 3) provide data for species assessments, management responses, and revision of the 23-year old Species Recovery Plan (Phillips and Phillips 1987). 2010] VI, ots rad Nee Y V5 _ === Humphreys Peak Trail mom \/Veatherford Trail _ == 200m Contour WHA; Estimated Packera franciscana Distribution Fic. 2. FOWLER AND SIEG: PACKERA FRANCISCANA DENSITY i) N ae | fe _1:24,000 ~ 0) 0.35 0.7 1.4 Kilometers [ee ee Site map showing the approximate distribution of Packera franciscana and the location of sampling transects along recreational trails on the San Francisco Peaks. The Humphreys Peak Trail transect is on the outer volcanic slopes through and above treeline, and the Weatherford Trail transect is mostly within the inner caldera basin but crosses to the outer slopes near its junction with the Humphreys Peak Trail. Distribution map was modified from Dexter (2007) to group his polygons and include areas where we found P. franciscana. Packera franciscana was not found at the top of Humphreys or Agassiz Peaks but was found at the top of Fremont Peak. METHODS In 2008 we established an elevational transect along the Humphreys Peak Trail on outer slopes of the San Francisco Peaks through P. francis- cana habitat to estimate the density of P. franciscana ramets, mid-September flowering/ fruiting phenology, and the population centroid elevation as the population intersects the trail (Fig. 2). Fifty seven sample points were estab- lished at 25 m intervals along a transect starting at 3520 m elevation, below the first P. franciscana occurrence, and extending 1425 m along the trail to an elevation of 3798 m which is above the last trail-side occurrence. In mid-September 2008 and 2009 after the monsoon season, we counted P. franciscana ramets (upright stems) within 12 individual | m° frames at each sample point arranged along the trail edge to allow flexibility for trail curvature (Fig. 3). Ramet counts were chosen as an estimate of abundance since P. franciscana 1s a rhizomatous species (Barkley 216 ___ YP Trail Left3 Right 3 Up Trail _— | Left 4 rege Left2 ~—-Right 2 _ Left 3 Right4 ~~ ‘Left2 Right1 1m | Let 4 ey Left1 Right 4 __|Left 1). Right2) | 1m im : | Recorded points oe ‘Left 6 Right 5 | | nee Down Trail Humphreys Peak Trail 1m Left 5 Right 5 Left 6 Right 6 ~ Down Trail — Weatherford Trail Fic. 3. Sampling frame layout along the Humphreys Peak and Weatherford Trails transect showing the 0-— 1 m and 1-2 m bands. Both layouts had 12 sampling frames per sample point. 1968; Trock 2006). Sampling frames were omitted when they overlapped previously counted frames along trail switchbacks, covered recent trail maintenance areas, or covered vertical drop-offs >5 m. Counts of ramets with flower, fruit, or both were also made within each frame. In September 2009 we added an inner basin transect along the Weatherford Trail (Fig. 2) to increase sample size and habitat diversity. This hiking trail runs mostly within the San Francisco Peaks volcanic caldera and crosses a large talus slope. This transect began at 3449 m, below where the first P. franciscana plants were noted, and continued with 110 sample points at 25 m intervals along the trail to its junction with the first trail at 3569 m. Sampling intensity, 12 frames per sample point, was the same as that on the first trail but arrangement of the sampling frames was restricted to within one meter of the trail edge due to the large amounts of loose talus off trail (Fig. 3). Coordinates for latitude, longitude, and elevation were made for each sample point with a Trimble® Geo XT 2005 Series GPS (Trimble Navigation Ltd, Sunnyvale, CA) with sub-meter accuracy to allow relocation of each sample point in subsequent years. Descriptive and inferential statistics were calculated with SAS/STAT 9.2 (SAS Institute MADRONO [Vol. 57 Inc, Cary, NC). Population centroid elevations were calculated in Proc Means as the elevation of each sample point weighted by the number of ramets, so that each ramet received equal weight. Proc GLIMMIX with a negative binomial distribution function was used to test for type III fixed effects differences in ramet count between years and trail-side bands (distance from trail; O-1 m vs. 1-2 m). Proc GLIMMIX with a normal distribution function was used to test for differences in elevation of P. franciscana occur- rences between the inner basin and outer slopes. The Standardized Morisita index of dispersion was calculated to measure ramet count aggrega- tion (Krebs 1989). RESULTS Packera franciscana occurred from 3529 to 3722 m elevation within two meters of the Humphreys Peak Trail transect. Within its range of occurrence along the trail, densities within the two-meter bands along each side for 2008 and 2009 were 4.18 and 2.74 ramets m ’, respectively (Table 1). There was no significant difference in overall ramet density between years (Fy. 0.05 = 0.45, P = 0.502). There was also no significant difference in ramet density between the 0-1 m bands and the 1—2 m bands (F). 9.95 = 2.43, P = 0.123) for either year (Fy. 9.95 = 0.34, P = 0.561). The elevation of the P. franciscana population centroid along the Humphreys Peak Trail, weighted by ramet density at each sample point, was 3640 (+5 SE) m for 2008 and 3641 (+7 SE) m in 2009 (Table 1). The overlapping standard errors indicate no significant change in elevation between years. In 2009, we added transect sampling points for P. franciscana along the Weatherford Trail, using only the 0-1 m band due to the large amount of loose volcanic talus just off the trail. Packera franciscana was found along this trail from 3471 m elevation to its junction with the Humphreys Peak Trail, both on the inner basin side and the outer volcanic slopes. In September 2009 we counted 1315 ramets of Packera franciscana at 163 sample points along the two transects. Ramet counts per sampling frame ranged from 0 to 180 and reflected the species’ visual patchiness. The Standardized Morisita index of dispersion (I, = 0.54) indicates a clumped pattern of ramet counts since it 1s between 0 (random) and +1 (maximum aggrega- tion). The elevation of the 2009 population centroid for both transects combined was at 3586 (+10 SE) m (Table 1), but the inner basin centroid, 3547 m, was significantly lower than the outer slopes centroid, 3638 m (F(o.05) 1, 28 = 14.92, P < 0.001). The overall P. franciscana 2009 trailside at) density estimate was 4.36 ramets m ~* within its 2010] FOWLER AND SIEG: PACKERA FRANCISCANA DENSITY 217 TABLE 1. ESTIMATES FOR DENSITY AND ELEVATION OF POPULATION CENTROIDS FOR PACKERA FRANCISCANA ON THE SAN FRANCISCO PEAKS IN NORTHERN ARIZONA. Density estimates follow a negative binomial distribution in which variance (var) is described by the negative binomial dispersion factor (k) and the square of the mean. The value of “‘k” given here was estimated by Proc GLIMMIX during statistical comparisons of the respective density estimates; variance in this table is the sample variance. Estimates for elevation of the population centroid are the mean of sample point elevations weighted by P. franciscana density which follow a normal distribution with standard errors (SE). Density ramets Sample location m ~ var Humphreys Trail 2008 4.18 2.27 0-1 m 2,02 31.65 1-2 m 6.41 294.13 Humphreys Trail 2009 2.74 44.58 0-1 m Ded 40.66 1-2 m 3.3] 71.82 Inner basin 2009 5.62 143.86 Outer volcanic slopes 2009 2.89 48.57 Overall 2009 4.36 101.58 elevational range of occurrence, 3471—3722 m on the San Francisco Peaks. The 2009 inner basin density was higher, 5.62 ramets m-*, than the estimate for the outer slopes, 2.89 ramets m * but not significantly SO (Fvo.05, 1,113) — 2.82 ,P= 0.096). The number of P. franciscana ramets within two meters of the Humphreys Peak Trail and within one meter of the Weatherford Trail is over 36,000 (density estimate * sampled length). Phenological measurements for P. franciscana during our mid-September sampling period were similar for 2008 and 2009, with 9% of the ramets either flowering and/or fruiting in 2008 versus 6% in 2009. There was less than 1% difference between inner basin and outer slopes flowering/ fruiting rate in 2009. DISCUSSION The Recovery Plan (Phillips and Phillips 1987) offers an overall estimate of 100,000+ clones of P. franciscana on the San Francisco Peaks as a general estimate of population size. Phillips and Peterson (1980) reported a P. franciscana popu- lation density range of 50-370 plants 100 m * on Agassiz Peak near the Weatherford Trail but did not clearly define plants as ramets or genets (clumps or clones) or describe estimation tech- niques. However, later references to clump size would indicate that they were using the genet concept. On a per 100 m° basis, our density estimate (436) is somewhat larger than the upper end of their density range (50-370), which may reflect the different “‘plant’’ definitions. Given the difficulty of defining and counting clumps or clones in the field, ramets provide a more accurate way to assess population density. Even though ramet density may inflate the number of functional plants, it is an accurate reflection of photosynthetic and reproductive potential. Phil- lips and Peterson (1980) also reported that 13% Centroid n k elevation SE 42 7.9 3640 m 5 im 42 9.7 42 Oey. 42 7.9 3641 m =) mM 42 9.7 42 9.7 a. 9.6 3547 m +7 TM 63 9.6 3638 m +7 m 141 na 3586 m = 10 i of the P. franciscana plants were adult (sexually reproducing) which is comparable to the 6—9% of ramets we sampled which were flowering and/or fruiting in 2008—2009. Our results are consistent with the above data from the 1980’s and give no indication of changing populations trends. AI- though these trail-side transects do not represent randomly selected population transects, they may be the only viable option since P. franciscana can inhabit large talus slopes which are very difficult to sample without uprooting plants near and within the sampling frame. These transects do sample the range of occupied habitats and observed densities in the center of its distribution in the San Francisco Peaks (Fig. 2). Our results and the estimate of >36,000 P. franciscana ramets in <2% of its range would indicate that the species is locally abundant, persisting, and reproducing. We interpret the successful production of fruit, which we observed actively dispersing by upslope winds in mid-September 2008, as an indication that P. franciscana can sexually reproduce on the San Francisco Peaks. Seed viability studies may provide additional support for this interpretation. Examination of plant root systems would be necessary to determine if new ramets originate from seed or from existing perennial rhizomatous clones. The hypothesis that rhizomes produce large patches of ramets is supported by the clumped pattern of ramet counts (I, = 0.54). Although this may be the primary method of reproduction (USFWS 1983), we also found single isolated ramets during our sampling which could be the result of seed dispersal or rhizome fragments moving downslope in the talus sub- strate that P. franciscana inhabits. Plants inhab- iting the upper portions of talus slopes would seem to be the result of seed dispersal since avalanches and downslope creep of talus fields would carry existing P. franciscana plants down- MADRONO [Vol. 57 Fic. 4. Photo of Packera franciscana (Greene) W.A. Weber and A. Love herbarium specimen showing ramets from an extensive rhizome and adventitious root system. slope. We noted dead P. franciscana plants at the base of some avalanche chutes. Our observations during voucher specimen collection indicate a relatively large root system comprised of rhi- zomes and adventitious roots that may not be attached to a stable substrate (Fig. 4). This growth habit in an unstable talus sea may be an evolutionary adaptation for survival and repro- duction in that fragmentation of the rhizome by talus creep processes may be common. Thus P. franciscana may be well adapted to this type of disturbance. The overall population centroid of 3586 m we measured is within the 3525-3605 m elevation range for most P. franciscana noted by Dexter (2007) and the 3350-3750 m main occurrence range in earlier reports (Phillips and Peterson 1980; U.S. Fish and Wildlife Service 1983). However, the population centroid for the out- slope samples located on a dry west-southwest slopes is 91 m higher in elevation than for the more east facing inner basin samples. The fact that southwest slopes have a higher P. franciscana patch centroid elevation lends credence to an upward migration hypothesis for this species in a future warmer drier climate. We plan annual measurements of both tran- sects to detect P. franciscana population trends. Sampling in subsequent years may indicate trends in population density, changes in September phenology, or elevational migration within its habitat. Changes in population density over time may allow detection of climate change effects, population cycles, or recreational impacts. Changes in the elevation of population centroids or September phenology will more likely be the result of climate change. ACKNOWLEDGMENTS Thanks to Brian Casavant and Addie Hite for help with fieldwork, sampling protocols, and dedication to this research. 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FROMENTIN, O. HOEGH-GULDBERG, AND F. BAIRLEIN. 2002. Ecological responses to recent climate change. Nature 416:389—395. MADRONO, Vol. 57, No. 4, pp. 220-228, 2010 PLANT POPULATION AND HABITAT CHARACTERISTICS OF THE ENDEMIC SONORAN DESERT CACTUS PENIOCEREUS STRIATUS IN ORGAN PIPE CACTUS NATIONAL MONUMENT, ARIZONA GRETA ANDERSON School of Geography and Development, University of Arizona, P.O. Box 210076, Tucson, AZ 85721 SUE RUTMAN National Park Service, Organ Pipe Cactus National Monument, 10 Organ Pipe Drive, Ajo, AZ 85321 SETH M. MUNSON! U.S. Geological Survey, Southwest Biological Science Center, Canyonlands Research Station, 2290 S. West Resource Blvd., Moab, UT 84532 ABSTRACT Peniocereus striatus (Brandegee) Buxb. (Cactaceae) is an endemic Sonoran Desert cactus that reaches its northern range limit in southwestern Arizona. One U.S. population occupies a small area of Organ Pipe Cactus National Monument near the U.S./Mexico international boundary, which has been monitored since 1939. An extensive survey conducted in 2002, covering 177 ha, resulted in the discovery of 88 new plants, in addition to the relocation of 57 plants found in previous surveys. Despite potential increases in population size and spatial distribution, mean plant height and number of basal stems has not significantly changed in recent years. Bud scars revealed that a majority of the population was sexually mature. Peniocereus striatus occurrence increased with decreasing slope, spanned every slope aspect, and was highest on rocky soils, but was noticeably low on west and northwest slopes and areas where severe land degradation had previously occurred. Over half of P. striatus plants were nursed by shrubs and subshrubs, while 40% occurred under leguminous trees. A severe frost in January 2002 top-killed 19% of the population, with the greatest damage in drainage bottoms. However, long-term (1944-2002) climate records show that there has been an overall increase in the number of frost free days in the region, which, coupled with land use change, has implications for the future health of this population. Key Words: Cardoncillo, climate, frost-tolerance, gearstem cactus, habitat suitability, land use history, night-blooming cereus, nurse plant. The small population sizes, narrow geographic ranges, and high habitat specificity make rare endemic plant species particularly vulnerable to accelerated climate and land use changes (Rabin- owitz 1981; Malcolm et al. 2006). The viability and persistence of rare endemic plants depends on the maintenance of suitable habitat and the ability of the population to propagate itself under changing environmental conditions. In the So- noran Desert, several endemic plants, including cacti, are of tropical descent and known to be limited in their northern distribution by freezing temperatures (Shreve 1911; Hastings 1963). Anthropogenic global warming is likely to decrease the intensity and frequency of freezing temperatures, which may create opportunity for expansion of frost-intolerant plant species (Ly- ford et al. 2003). In addition to climate, Sonoran Desert endemic plants are likely to respond to changes in land use that occur in their local ' Corresponding author: SMunson@usgs.gov habitat, including livestock grazing, wood har- vesting, cropping practices, and other activities (Suzan et al. 1994). Close study and monitoring of plant populations and their habitat at the periphery of their geographic ranges can contrib- ute to an understanding of the factors that limit their distribution, provide information on the sustainability of the population, and help inform conservation strategies. Peniocereus striatus (Brandegee) Buxb. is a slender-stemmed cactus endemic to frost-free areas of the Sonoran Desert (Felger 2000). This inconspicuous sub-erect to sprawling, vine-like | cactus 1s typically 25—75 cm tall (Pinkava 1995) and up to 2 m tall in some microhabitats (Felger | 2000). Peniocereus striatus has small, scattered populations throughout Mexico (Baja California, , Sonora, and Sinaloa) and is very rare in the U.S., | where it reaches its northern range limit in | southwestern Arizona (Benson 1982; Nabhan | 1992; and Felger 2000). One of the few P. striatus ' populations in the U.S. occurs in a small area of | Organ Pipe Cactus National Monument | 2010] (OPCNM) near the U.S./Mexico international boundary. This population was first recorded from Gray’s Well in 1939 (OPCNM Herbarium, NE of Sonoyta, 20 April 1939, A.A. Nichol s.n.), and no other population has been reported from any other area in OPCNM. Past research has found that populations of P. striatus have higher densities and larger plant sizes south of OPCNM at less disturbed sites (Suzan et al. 1994). The condition of P. striatus improves under perennial nurse plants, which modify the environment by buffering extreme temperatures, increasing water and nitrogen availability, reducing photosynthetically active radiation, providing physical support and pro- tection from herbivores (Franco and Nobel 1989; Suzan et al. 1996). Recruitment of P. striatus 1s low, most likely because it has few species of pollinators, blooms nocturnally during a few episodes, and has low floral and fruit production (Benson 1950; Suzan et al. 1994; Raguso et al. 2003). Previous reports suggested that P. striatus in OPCNM occurred more frequently on rocky hill slope habitat (Jackson 1966; Steenbergh 1966; Suzan et al. 1994; Goodsell unpublished), which contrasts with observations that P. striatus in Mexico prefers flat sites more than rocky slopes (T. Van Devender and M. Dimmitt, Arizona- Sonora Desert Museum, personal communica- tion; G. Anderson and S. Rutman personal observation). We hypothesize that P. striatus occurred on rocky hill slopes in OPCNM during a historical period of high land use intensity because these sites provided refugia from plant mortality and habitat degradation associated with livestock grazing and other land manage- ment practices. We predict that the population of P. striatus in OPCNM has expanded from the area occupied during the original survey for two main reasons. First, deleterious management practices have ceased, potentially allowing for increased recruitment, germination, and survival of P. striatus plants, as well as the regeneration of P. striatus nurse plants, which create suitable habitat for growth. Second, the frequency and duration of freezing temperatures in the winter has likely decreased since the population was first recorded. As winter freezing temperatures de- crease, the survival and reproduction of these frost-intolerant plants might increase. In this study, we use over 60 years of monitoring results to determine the population and habitat characteristics of P. striatus in OPCNM and provide some evidence of how climate and land use changes can affect succulent plants at the limits of their distributions. Our specific objectives were to: 1) determine the size, spatial distribution, and reproductive capacity of the population; 2) identify suitable habitat characteristics, including the use of nurse plants; ANDERSON ET AL.: CHARACTERISTICS OF PENIOCEREUS STRIATUS IN OPCNM 221 and 3) assess past and current climate and land use threats to the population METHODS Site Description Organ Pipe Cactus National Monument is an International Biosphere Reserve located in south- western Arizona, 210 km W of Tucson and 35 km S of Why, AZ (31.82—32.20°N, 112.61-— 113.09°W). Organ Pipe Cactus National Monu- ment includes several high mountain ranges and low basins, with elevations ranging from 300— 1470 m. Long-term (1944-2002) mean annual temperature at low elevation is 21.0°C (mean min. = 12.1°C, mean max. = 29.9°C) and mean annual precipitation (MAP) is 238 mm. Precip- itation at OPCNM is bimodally distributed, with nearly half of the precipitation delivered during winter (October—March) storms and the remain- der during the summer monsoon (July—Septem- ber). April-June are extremely dry, with total precipitation in these months composing 4% of MAP. Organ Pipe Cactus National Monument was established in 1937 to protect the organ pipe cactus (Stenocereus thurberi (Engelm.) Buxb.) and the Sonoran Desert ecosystem. The OPCNM Peniocereus striatus population occurs within a Larrea tridentata-Ambrosia deltoidea association on floodplains and lower bajadas, an Atriplex polycarpa-A. linearis association on loamy floodplains, and a Prosopis velutina- Parkinsonia microphylla-Olneya_ tesota associa- tion on hill slopes. Monitoring History of the P. striatus Population in OPCNM Thirty years following the first documentation of P. striatus in OPCNM in 1939, the National Park Service (NPS) reported the species was rare and occurred only on north- and east-facing hill slopes (Jackson 1966; Steenbergh 1966). In 1969, the (NPS) surveyed in the proximity of the original location and reported a total population of 36 plants on three hills (Goodsell unpublished). As part of the Sensitive Ecosystem Program in OPCNM, probable habitat, defined as hill slopes in the vicinity of the plants found in 1969, was surveyed for P. striatus in 1990. The survey resulted in the location of 59 plants (11 plants per ha in suitable habitat), which produced an estimate of the extant population in OPCNM of less than two hundred individuals (Johnson et al. 1990; Ruffner and Associates 1995). Asa result of this survey, the NPS measured the height and survivorship of 22 P. striatus plants from 1991— 1996. Another survey occurred in 1994, when 47 plants in OPCNM were located and tagged for a 222 study on the pollination ecology of P. striatus (Nabhan and Suzan 1994). Some of these individuals were relocated in 1999, when volun- teers surveyed the eastern portion of its range in OPCNM. Fifty-seven plants were found, tagged and mapped. About two dozen P. striatus plants were tagged in 1991 and revisited from 1993-1999 to determine population characteris- tics (Ruffner and Associates 1995), with little to no information recorded about habitat require- ments. 2002 Survey To determine the population size, distribution, and habitat preferences of P. striatus in OPCNM, a systematic survey was conducted from Janu- ary—July 2002. Most of the survey was performed by a single person trained in the appearance and growth habits of the plant. Survey intensity ranged from belt transects (S—10 m wide) in low density vegetation to more intensive surveys in xeroriparian areas. All open areas and every nurse plant species was checked for the presence of P. striatus. The survey was initiated in previously occupied habitat determined by previ- ous NPS reports (Jackson 1966; Steenbergh 1966; Goodsell unpublished) and expanded into poten- tial habitat until the survey no longer located plants or the international boundary was encoun- tered. Survey tracks were recorded with a high precision GPS unit (accuracy ~3 m), and survey area was created by buffering survey tracks in a GIS. The location and elevation of P. striatus plants were also recorded using a high precision GPS unit. Plants were marked with numbered metal tags and the maximum height and width of each plant, as well as the number of live and dead basal stems per plant were recorded. Height and width measurements were taken during winter dormancy and only green succulent stems of plants were measured. To determine herbivory damage, the presence or absence of injury to a plant and proximity to a rodent hole were recorded. Since flower bud scars are obvious and persist for a long time, they were used to indicate reproductive status. The nurse plant (sensu Franco and Nobel 1989) and its spatial orientation to P. striatus, as well the nearest perennial plant neighbor and its distance to P. striatus were noted. Slope, aspect and other descriptors of the physical environment were recorded. Soil surveys from OPCNM (USDA-Soil Conservation Ser- vice 1972) were used to produce a digital data layer of soil types within the P. striatus survey area. Lastly, whether or not the plant was within two meters of an ephemeral drainage was observed and recorded. MADRONO [Vol. 57 Climate and Land Use History Precipitation was measured at the Dos Lomitas rain gauge, located within the study area, and temperature was measured at the National Weather Service station located at park head- quarters 20 km NW of Dos Lomitas. To assess past land use history of the study area, a map of historic land use was prepared. A digital ortho- photo quadrangle, formed from a 1997 photo- graphic image, was used as a base layer onto which lines and polygons were drawn to indicate roads, corrals, wells, historic agricultural fields and disturbance zones. Early aerial photographs of the area were compared with the 1997 ortho- photo quad to verify or interpret linear features and other patterns of land use. Some man-made features found on the ortho-photo quad were verified on the ground using GPS. Archived documents at OPCNM provided additional information on past land management activities (Rutman 1996). Statistical Analyses The height and width of P. striatus in 2002 were broken into 10 cm size classes to assess population structure. Changes in height and number of basal stems through time were evaluated using repeated measures ANOVA (R Development Core Team 2008). The proportion of the total population in each slope class, aspect, soil type, nurse plant, and nearest perennial plant was determined. Pearson’s Chi-square tests were performed to determine if observed occurrences were different than an expected random distribu- tion in potential habitat. ANOVA was also performed on height of P. striatus to see if there were differences among nurse plant associations. A linear regression model was fit to determine trends in temperature through time. RESULTS AND DISCUSSION Population Characteristics A survey of 172 ha of potential Peniocereus striatus habitat in OPCNM resulted in _ the discovery of 88 new plants, in addition to the relocation of 57 plants found in the 1999 survey, which makes the density of the population 0.85 individuals per ha. This population size of 145 plants is larger and population density two orders of magnitude greater than estimates reported by Suzan et al. (1994). The close proximity (<3 m) of many new P. striatus found in the 2002 survey to those found in 1969 (Fig. 1), and the discovery of small individuals in 2002, provide further evidence that there have likely been increases in the population size. The population also appears © to have expanded from its restricted spatial | 2010] : a wit re < EY. © wens é 0.4 Miles 0.8 Kilometers Peniocereus striatus plant, 1969 ake Peniocereus striatus plant, 2002 2002 Survey perimeter BiG. J Organ Pipe Cactus National Monument. distribution on hillslopes in 1969 to adjacent low- lying areas, but individuals were notably absent from the farthest western portion of the survey area where intense land use modification had occurred (Fig. 1). Potential increases in popula- tion size and spatial distribution may be partially due to differences in survey efforts, which were not well described in historic records. The mean height of plants in 1993 did not significantly change in subsequent surveys in 1994-2002 (Fig. 2a). The mean number of basal stems did not significantly change from 1999 to 2002, although there was a trend for stem number to increase (Fig. 2b). The mean height and width of plants were 58.6 cm (+2.5 cm standard error) and 54.4 cm (+3.7 cm), respectively. The height Dike ANDERSON ET AL.: CHARACTERISTICS OF PENIOCEREUS STRIATUS IN OPCNM 225 Organ Pipe Cactus National Monument _»..... Py a Bs Pe Scale = 1:15,000 UTM12N NAD83 i June 2010 Erosion control structure a | Organ Pipe Cactus NM boundary Location of Peniocereus striatus individuals from 1969 and 2002 surveys and land use disturbances in distribution of the population ranged from 1 to 158 cm and had a significant (>2 standard errors of skewness) positive skew (1.02), which means the tail of the distribution was shifted towards taller individuals (Fig. 3a). While a majority of individuals in the P. striatus population were 40 80 cm tall, it is likely that the interaction P. striatus had with nurse plants allowed for the growth of tall individuals by facilitating an environment that was favorable for plant growth. The tallest plant of 158 cm was close to the maximum recorded height of 200 cm (Felger 2000; Goodsell unpublished). The horizontal width distribution was bimodal, with most individuals <10 cm or 30-70 cm (Fig. 3b). This reflects the vine-like (narrow) and bush type (wide) morphological types first described by Johnson et al. (1990). The vine-like morphs often occurred under trees and had long branches that draped across nurse plant stems, while the bush- type morphs appeared to have shorter, more erect branches and usually grew within the canopies of shrubs and subshrubs or in the open. The maximum width of 304 cm exceeded previously reported values (Johnson et al. 1990; Pinkava 1995; Anderson 2002). Most plants had a single living basal stem, although at least one plant had eight stems. A majority of P. striatus individuals in the OPCNM population were sexually mature. The presence of bud scars indicated that 76% of the plants had flowered in the past. For the remainder of the population, reproductive status could not be determined, either due to herbivory or aboveground tissue damage. In July 2002, 45% of P. striatus individuals had buds _ present, including two plants that lacked bud scars from prior reproductive activity. Habitat Peniocereus striatus occurred at a low, fairly narrow elevation range between 388-441 m, which is within the elevation range of 60-450 m reported for this species (Benson 1982). The number of plants decreased with increasing slope: 35% of plants occurred on <2% slope, while only 5% of plants occurred on >22% slope (Table 1). This suggests that there has been expansion of the population from its restricted historical hill slope habitat. Peniocereus striatus occurred on every slope aspect, but had a tendency to occur less frequently on west and northwest facing slopes (x? = 21.5, P = 0.003, 224 MADRONO [Vol. 57 (a) (a) 20 1 ee 5 60 = wa fe) — = 15 = 50 a oh * za v iS) 10 4 O40 es i) x 5] a (b) 5 30 al fae 2 4 nN BPW HD OPH PG GP HW HG. a SNAG ICUS vy” ery” 7 ge 25 \ POP PS SN HL WG Loyrnyr wry a ai (b) Height Class (cm) oa. a: 20 4 io) a “ites 5 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 = i 3 Y ear S. < 10 Fic. 2. Mean height (a;n = 15) and total number of ‘% basal stems per plant (b; n = 51) of Peniocereus striatus Snes from 1993-2002. No significant differences among years (P > 0.05). 1 PO HPP OPS opt Newsn ow PoP WIS MH wigs Se et Ca 7 Width Class (cm) Fic. 3. Percent of the Peniocereus striatus population in height (a) and width (b) classes. Height distribution skewness = 1.024, kurtosis = —0.375. Skewness and kurtosis is considered significant if greater than 2 standard errors of normal distribution = +0.436. df = 7; Fig. 4a). This result is consistent with earlier reports that suggested P. striatus occurred less frequently on west-facing slopes (Steenbergh 1966; Goodsell unpublished). While many cacti in OPCNM attain their greatest densities on south- facing slopes since they are less subject to freezing temperatures in the winter (Parker 1987), the current P. striatus distribution is not limited to these slopes, perhaps because they are frequently buffered from extreme environmental conditions by nurse plants. Peniocerius striatus occurred on five different soils and three landform types within the 2002 survey area (Table 1). Plants occurred less frequently on fine and very fine sandy loams and more frequently on rocky soils than predict- ed if plants were randomly distributed in their potential habitat (y° = 109.4, P < 0.0001, df = 5). Twenty-seven percent of the population occurred on deep, very fine or fine sandy loam on gently sloping floodplains, nearly half of the plants occurred on very gravelly or cobbly loam on dissected alluvial fans, and another quarter of the plants grew on very stony loam on hill slopes (Table 1). Peniocerius striatus was not found on torrifluvents of the drainage bottoms, although this substrate type only represented 4.3 percent of | the survey area. Although these results are consistent with historical accounts of greater P. 2010) ANDERSON ET AL.: CHARACTERISTICS OF PENIOCEREUS STRIATUS IN OPCNM 225 TABLE 1. HABITAT CHARACTERISTICS OF PENIO- (a) CEREUS STRIATUS POPULATION AT ORGAN PIPE CACTUS NATIONAL MONUMENT. N 15% Habitat % of population NW ie NE Slope 0-2% 35 2% 3-8 % 25 a ; 9-14% pA 15-21% 14 >22% 5 Soil type SW SE Gunsight very gravelly loam 4] : No Aspect =21% Gilman very fine sandy loam 26 : Lomitas very stony loam 24 Harqua very cobbly loam 8 (b) Antho fine sandy loam l Torrifluvents 0 striatus occurrence on rocky soils, the population has expanded to other soil types. Nurse Plant Associations Most P. striatus plants in OPCNM grew within the canopy of subshrubs, shrubs and trees. Nabhan and Suzan (1994) also found that P. striatus was non-randomly distributed in the environment, preferring microsites under nurse plants to microsites with no plant cover. Over half of P. striatus plants were nursed by shrubs and subshrubs, while 40% occurred under leguminous trees (Table 2). These results are similar to Nabhan and Suzan (1994), who found that 50% of P. striatus grew under trees. A majority of P. striatus that had nurse plants were directly within them, while most others were oriented either directly north or south of the nurse plant (y° = 47.0, P < 0.0001, df = 7; Fig. 4b). An orientation north of the nurse plant can be explained by protection against excessive radiation, temperature, and water stress (Franco and Nobel 1989), while an orientation south of the nurse plant may provide protection against frost (Suzan 1994). While we found Prosopis velutina Wooton to be the most commonly used nurse plant by P. striatus, Nabhan and Suzan (1994) reported Olneya tesota A. Gray was the species’ principal nurse plant. Besides providing shade and physical protection, legume trees also increase soil nitro- gen levels under their canopies (Franco and Nobel 1989). Johnson et al. (1990) observed that Ambrosia deltoidea (Torr.) W. W. Payne, Larrea tridentata (DC.) Coville, and Parkinsonia micro- phylla Torr. were the most common nurse plants for P. striatus. The earliest report of the OPCNM population mentions Larrea tridentata as the most common associate (Goodsell unpublished), followed in quantity by plants standing entirely alone. Nurse plant type significantly affected the NW NE SE Directly under nurse plant = 23% SW S Fic. 4. Percent of the Peniocereus striatus population on each aspect (a) and orientation in relationship to nurse plant (b). Aspect: y° = 21.4, P = 0.003, df = 7, orientation to nurse plant: y° = 50.2, P < 0.0001, df = 7. Chi-square test assumes expected population is evenly distributed in each direction. mean height of P. striatus. Plants underneath trees were significantly taller than those under subshrubs, while plants underneath shrubs and those with no nurse plant association had an TABLE 2. NURSE PLANT ASSOCIATIONS OF PENIO- CEREUS STRIATUS POPULATION AT ORGAN PIPE CACTUS NATIONAL MONUMENT. Nurse plants % of population Subshrubs 25 Atriplex linearis S. Watson IB Ambrosia deltoidea (Torr.) W. W. Payne [2 Shrubs 29 Larrea tridentata (DC.) Coville 14 Atriplex polycarpa (Torr.) S. Watson 1] Lycium spp. 4 Trees 40 Prosopis veluntina Wooton 21 Parkinsonia microphylla Torr. 12 Olneya tesota A. Gray 7 Other 6 None 4 Prosopis veluntina (dead) l Unknown shrub (dead) l 80 60 40 0 Mean Height of P. striatus (cm) Tree Subshrub Shrub Nurse Plant Fic. 5. Mean height of Peniocereus striatus in relationship to the nurse plant type (none, subshrub, shrub, tree). Different letters designate significant differences (Tukey adjusted P < 0.05). None intermediate height (Fig. 5). This supports our hypothesis that P. striatus makes opportunistic use of the structural support of the available perennial canopy and that the shrubby morph is more likely to be found in the open or under a subshrub. Peniocereus striatus was on average 16.7 (+1.7) cm from the nearest perennial plant, which was most commonly a subshrub (55%), either Ambro- sia deltoidea or Atriplex linearis S. Watson, followed in close proximity to shrubs (30%). Threats to Population Climate. Most plants had at least one dead basal stem, indicating that aboveground tissue 40 5 i = MADRONO [Vol. 57 damage was widespread and common. During the winter-spring survey of 2002, 19% of P. striatus individuals with at least one previously green stem had all their stems become light brown, dry and brittle. Since most of the plant’s biomass is belowground in diffuse tubercles (Benson 1982), this “‘top-kill’’ does not necessar- ily equate to plant mortality. One likely explana- tion for this observed “‘top-kill” is freezing temperatures. Temperature at the site dropped to —13.7°C on January 31, 2002, which is low enough to cause tissue damage or death of temperature-sensitive succulents (Nobel 1988). Position on the landscape significantly affected the probability of the 2002 aboveground tissue damage (y* = 70.4, P < 0.0001, df = 3): 36% of individuals that occurred =2 m from a drainage showed top-kill damage while only 8% of individuals >2 m from a drainage were damaged. Since drainages are frequently affected by cold nocturnal air inversions due to radiative cooling and cold air flow from sidewalls (Brunt 1939), the higher incidence of top-kill in low-lying areas was likely caused by low temperatures. By early summer 2002, more than half of top-killed plants had re-sprouted from the base or had new branches growing from the desiccated stems. The likelihood of frost damage is likely to decrease in the future as temperatures warm due to anthropogenic greenhouse gas emissions. Over the last fifty years (1949-2002), the number of freezing days has declined from greater than 20 to less than 10 days per year (linear regression: slope = —0.33, r? = 0.41, P < 0.0001; Fig. 6), the average minimum January (coldest month in OPCNM) temperature has increased 2.3°C, and this average has not been below freezing since rer o’. | | 6 ‘A L | T T T T T T L980 L985 1990 L998 2000 2005 T 1970 1975 Year Annual number of freezing days at study site from 1949-2002 (black line) and 10-year moving average 0.41, P < 0.0001. T T T T LOSS 1960 1965 Qos L9AS Annual number of freezing davs ee Cc 1950 PIG: G. (gray line). Linear regression: slope = —0.33, r? 2010] 1949 (not shown). Given that minimum temper- atures likely limit the range of P. striatus, these changes in temperature may explain potential increases in the P. striatus population. Future increases in temperature are likely to result in the spread of frost-intolerant succulents and other plants northwards in their distribution. Herbivory. Most P. striatus plants (70% of the population) displayed signs of herbivore damage during the winter 2002 survey, likely caused by rodents and lagomorphs. In 2002, 17% of new stems initiated in the spring showed signs of herbivory by July. Nearly half (45%) of the total population was within | m of a rodent hole, which suggests a high animal density in close proximity to the cactus population. Land use. The habitat of P. striatus is located in an area of OPCNM heavily degraded by past land use and land management practices. Con- centrated livestock grazing, wood harvesting, and farming were among the land use activities within the study site at the Gray Ranch headquarters (Rutman 1996). The overstocking of cattle had adverse environmental effects, including damage to P. striatus and its habitat, which continued until livestock were removed in the late 1970's (BLM 1966; Schultz et al. 1971). On certain soil types in the study area, poor livestock grazing practices led to accelerated erosion, expressed as deeply entrenched channels and headcuts that moved upstream as much as 8 m per rainstorm in 1952 (OPCNM historic photo, accession #1629). Large stumps and re-sprouted stems at the study site are signs of historic tree harvesting, while more recently, illegal woodcutters have taken tree branches using machetes (Nabhan and Suzan 1994). Peniocereus striatus could not have been present on a 13 acre flood-irrigated field used by the Gray family at the north end of the study area. Small diversion dams and retention dikes used to manage irrigation, which date back to the early 1930’s, change water surface flows on several hundred acres of suitable habitat for P. striatus. Potential habitat in the study area has also been disrupted by erosion control structures installed by the NPS and Soil Conservation Service in the 1950’s—1960’s. More than 50 years after their construction, these structures remained clearly visible in aerial photography (northwest corer of Fig. 1). Despite these structures, erosion rates have not slowed since 1977 (T. Marsh, unpublished data, 1977-1996). The loss of soil has undoubtedly had consequences for the local P. striatus population, as the roots would have been exposed or buried, depending on landscape position. Peniocereus striatus has not been found in or adjacent to these erosion control structures, which suggests that the species may be sensitive to the local watershed modifications caused by these structures. ANDERSON ET AL.: CHARACTERISTICS OF PENIOCEREUS STRIATUS IN OPCNM 227 Previous surveys and knowledge about the study area’s land use history provides some evidence that the P. striatus population was historically restricted to rocky hill slope habitat, as reported by early surveyors. This landscape position was largely left undisturbed by livestock grazing, erosion, agriculture, and watershed modifications, which were concentrated on the low-lying and flatter portions of the landscape. The cessation of livestock grazing, coupled with warmer temperatures, may have allowed for the expansion of P. striatus into areas of the landscape where it was previously not found. Erosion control structures and their long-term effect on hydrology continue to exclude this species from otherwise suitable habitat. ACKNOWLEDGMENTS This work was supported by a grant from the USGS Status and Trends of Biological Resources Program. We wish to thank NPS and the Arizona-Sonora Desert Museum for their support and encouragement for this project. We are grateful to Tom Van Devender and Ana Lilia Reina, who showed G.A. Peniocereus striatus in Mexico, Peter Holm, who provided climate data, and Andy Hubbard for his input. Thanks to Kathleen Parker, Mary Moran, and two anonymous reviewers for helpful suggestions on how to improve earlier versions of this paper. Any use of trade, product, or firm names in this paper is for descriptive purposes only and does not imply endorsement by the U.S. Govern- ment. All authors contributed equally to this work. In memory of NPS Ranger Kris Eggle, 1973—2002. LITERATURE CITED ANDERSON, E. 2002. The cactus family. Timber Press, Portland, OR. BENSON, L. 1950. The cacti of Arizona, 2nd ed. University of New Mexico Press, Albuquerque, NM. . 1982. The cacti of the United States and Canada. Stanford University Press, Stanford, CA. BRUNT, D. 1939. Physical and dynamical meteorology, 2nd ed. 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Special 228 MADRONO status plants of Organ Pipe Cactus National Monument, AZ, Sensitive Ecosystems Project, Final Report. Organ Pipe Cactus National Mon- ument, Ajo, AZ. LYFORD, M. E., S. T. JACKSON, J. L. BETANCOURT, AND S. T. GRAY. 2003. Influence of landscape structure and climate variability on a Late Holo- cene plant migration. Ecological Monographs 73:567-583. MALCOLM, J. R., C. Liu, R. P. NEILSON, L. HANSEN, AND L. HANNAH. 2006. Global warming and extinctions of endemic species from biodiversity hotspots. Conservation Biology 20:538—548. NABHAN, G. P. 1992. Status survey of Peniocereus greggii and P. striatus in the Sonoran Desert. Contract report for U.S. Fish and Wildlife Service, Albuquerque, NM. AND H. SUZAN. 1994. Boundary effects on endangered cacti and their nurse plants in and near a Sonoran Desert biosphere reserve. Occasional Papers in Conservation Biology 1:55—67. NoBEL, P. S. 1988. Environmental biology of agaves and cacti. Cambridge University Press, Cambridge, U.K. PARKER, K. C. 1987. Site-related demographic patterns of organ pipe cactus populations in southern Arizona. Bulletin of the Torrey Botanical Club 114:149-155. PINKAVA, D. J. 1995. Cactaceae, pt. 1, The ceroid cacti. Journal of the Arizona-Nevada Academy of Science 29:9-10. R DEVELOPMENT CORE TEAM. 2008, R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria, Website: http://www.r-project.org/ [ac- cessed 12 December 2010]. RABINOWITZ, D. 1981. Seven forms of rarity. Pp. 205— 217 in H. Synge (ed.), The biological aspects of rare plant conservation. John Wiley and Sons, New York, NY. [Vol. 57 RAGusO, R. A., C. HENZEL, S. L. BUCHMANN, AND G. P. NABHAN. 2003. Trumpet flowers of the Sonoran Desert: floral biology of Peniocerius cacti and Sacred Datura. International Journal of Plant Science 164:877—892. RUFFNER AND ASSOCIATES. 1995. Organ Pipe Cactus National Monument ecological monitoring pro- gram monitoring protocol manual, Special Report No. 11, National Biological Service-Cooperative Park Studies Unit, University of Arizona, Tucson, AZ. RUTMAN, S. 1996. Livestock grazing and accelerated erosion on Organ Pipe Cactus National Monu- ment, Ajo, AZ. Organ Pipe Cactus National Monument, Ajo, AZ. SCHULTZ, R. L., E. L. WEBB, J. P. TOWNSEND, AND W. K. CARTER. 1971. Range conditions on the Organ Pipe Cactus National Monument-—a special study for the National Park Service. Bureau of Land Management, Arizona State Office, Phoenix, AZ. SHREVE, F. 1911. The influence of low temperatures on the distribution of the giant cactus. Plant World 14:136-146. STEENBERGH, W. F. 1966. Memorandum to the Superintendent, unpublished report, 1 May 1966. Organ Pipe Cactus National Monument, Ajo, AZ. SUZAN, H., G. P. NABHAN, AND D. T. PATTEN. 1994. Nurse plant and floral biology of a rare night blooming Cereus Peniocereus straitus (Brandegee) F. Buxbaum. Conservation Biology 8:461—470. : , AND . 1996. The importance of Olneya tesota as a nurse plant in the Sonoran Desert. Journal of Vegetation Science 7:635—644. U.S. DEPARTMENT OF AGRICULTURE (USDA)-SOIL CONSERVATION SERVICE. 1972, Soil survey—a special report: Organ Pipe Cactus National Mon- ument, Pima County, Arizona. Soil Conservation Service, Phoenix, AZ. MADRONO, Vol. 57, No. 4, pp. 229-241, 2010 STAND DEVELOPMENT ON A 127-YR CHRONOSEQUENCE OF NATURALLY REGENERATING SEQUOIA SEMPERVIRENS (TAXODIACEAE) FORESTS WILL RUSSELL AND KRISTIN HAGESETH MICHELS Department of Environmental Studies, San José State University, San José, CA, USA 95192-0115 will.russell@sjsu.edu ABSTRACT Understanding the natural patterns of regeneration following human disturbance is essential for effective restoration and management of second-growth forests. Despite their unique ecological character, little is known about these patterns in Sequoia sempervirens (D. Don) Endl. (Coast Redwood) forests. We examined the composition and structure of naturally regenerating stands with 360 randomly located sample plots across a chronosequence of five replicated age-classes (18 to 127 yr) and three old-growth reference sites. Results indicate a progression of stand characteristics towards old-growth conditions, with several measures reaching old-growth equivalence within the timeframe of the chronosequence. Stand density, canopy cover, and species richness reached old-growth equivalence within 41—80 yr; Shannon-diversity reached old-growth equivalence between 80—100 yr: and the density of redwood seedlings and shrub cover reached old-growth equivalence between 100— 130 yr. Basal area, herb cover, and the relative dominance of S. sempervirens progressed toward, but did not reach, old-growth equivalence. Size-class analysis indicated an increase in the density of large diameter trees, with no change in the density of smaller size-classes after forty yr. Coast redwood associated understory species were favored on the older sites with the cover of Calypso bulbosa (L.) Oakes, Trillium ovatum Pursh, and Viola sempervirens Greene reaching old-growth equivalence, while Tris douglasiana Herb., Tiarella trifoliate L., and Achlys triphylla (Sm.) DC. did not. No non-native species were recorded in stands older than 60 yr. We conclude that coast redwood forests are resilient to human disturbance, though some old-growth characteristics may require more than a century to develop. Key Words: Chronosequence, coast redwood, sempervirens. The coast redwood forest, dominated by Sequoia sempervirens (D. Don) Endl., is known for its high productivity, large carbon storage potential, and impressive stature (Preston 2007). Historically, S. sempervirens forests covered more than 8100 km? along the fog-shrouded coast from central California to southern Oregon. Due in large part to its value as a timber species, more than 95 percent of the original old-growth coast redwood forest has been converted into managed timber stands and other land uses (Noss 2000). With the majority of this forest type currently in second-growth, analysis of the natural patterns of stand development following timber-harvest is essential for effective management and restora- tion. The dynamics of post-harvest development for S. sempervirens stands are unique among conif- erous forests. Sequoia sempervirens possesses a natural resilience to disturbance due in part to its prolific vegetative sprouting ability (McBride 1977; Espinosa-Garcia and Langenheim 1991: Veirs 1996; Sawyer et al. 2000; Barbour et al. 200i). Though regeneration of S. sempervirens from seed can occasionally be abundant on mineral soils and fallen logs (Bingham and Sawyer 1988; Becking 1996; Porter 2002), the natural regeneration, second-growth, Sequoia majority of recruitment results from vegetative sprouts, especially following timber harvest (Douhovnikoff et al. 2004; Lorimer et al. 2009). The vascular connection between vegetative sprouts and existing root structures results in competition between stems for apical dominance, rather than for individual tree survival (Kauppi et al. 1987; Burrows 1990; Sachs et al. 1993; Laureysens et al. 2003). Issues of overcrowding that slow the regeneration of other coniferous trees do not affect S. sempervirens in the same manner. Instead, redundant clonal stems senesce over time, thinning the stand naturally without the risk of stand-scale mortality (Floyd et al. 2009; Lutz and Halpern 2006; Sach et al. 1993). Survival of suppressed trees is also unusually high for S. sempervirens as epicormic sprouting increases stem production when understory trees are released (Finney 1993). Old-growth S. sempervirens forests are rela- tively stable in terms of composition and structure (Busing and Fujimori 2002) and follow a ‘gap phase’ regenerative pattern where sup- pressed understory trees expand to fill canopy gaps created by individual, or small group, blow- downs (Sawyer et al. 2000). Natural stand- replacing disturbance events are extremely rare 230 in S. sempervirens forests, even when compared to other coniferous forest types in the Pacific Northwest (Lorimer et al. 2009). As a result, the stand-scale removal of canopy through timber-harvest initiates regenerative patterns oth- erwise undocumented in this forest type. Regeneration of S. sempervirens following timber harvest has been studied primarily in the context of post-harvest management practices such as planting, seeding, and thinning (Cole 1983; Oliver et al. 1994; Lindquist 2004a, b; Chittick and Keyes 2007; O’Hara et al. 2007). Very few studies have addressed the development of S. sempervirens stands in the absence of post- harvest management, and those that have been conducted focused on specific case studies or individual species rather than on forest regener- ation as a whole (Boe 1965; Powers and Wiant 1970; Allen et al. 1996; Jules and Rathcke 1999). Analysis of stand development of a long-lived species such as S. sempervirens (commonly exceeding 1500 yr in age) is best accomplished through the use of a chronosequence. This method has been routinely applied in other forest types (Crowell and Freedman 1994; Mund et al. 2002; Letcher and Chazdon 2009) as well as in S. sempervirens forests where they have been used to study specific impacts of logging (Loya and Jules 2007; Russell and Jones 2001; Russell 2009), but not over-all stand development. The objective of this study is to analyze natural regeneration of forest structure and composition in coast red- wood forests, with the hypothesis that stand characteristics will tend toward old-growth con- ditions over time. METHODS Study Sites Data was collected in the central range of the coast redwood forest, as defined by Sawyer et al. (2000) (Fig. 1). Study sites were located primar- ily in the Big River watershed, consisting of more than 2968 hectares of previously harvested coast redwood forest in Mendocino Co., Cali- fornia (California Department of Parks and Recreation 2006). Much of the watershed was managed as industrial timberland prior to its purchase by the Mendocino Land Trust in 2002, and subsequent transfer to the California State Parks as the Big River Unit of Mendocino Headlands State Park. The Big River watershed was an ideal location for this study due to the presence of second-growth redwood stands ranging from 15 to 127 yr old that had received no post-harvest manipulation. The vegetation of the area is characteristic of the central range of the coast redwood forest with Sequoia sempervirens, Pseudotsuga menziesii (Mirb.) Franco var. menziesii, and Abies grandis MADRONO [Vol. 57 (Douglas ex D. Don) Lindl. dominating the canopy, and Lithocarpus densiflorus (Hook. & Arn.) Rehder and Tuga heterophylla (Raf.) Sarg. commonly occurring in the subcanopy. Under- story shrub species include Polystichum munitum (Kaulf.) C. Presl, Vaccinium ovatum Pursh, and Rhododendron macrophyllum D. Don. Common herbaceous understory species include Oxalis oregana Nutt., Trillium ovatum Pursh, Viola sempervirens Greene, Calypso bulbosa (L.) Oakes, Tris douglasiana Herb., Tiarella trifoliata L., and Achlys triphylla (Sm.) DC. The soils of the area are derived from the Franciscan assemblage, consisting mainly of sandstone and marine sediments. Typically, winters are cool and wet with an annual precipitation of 2500 mm or more (Sawyer et al. 2000). Summers are mild with moisture from intermittent fog providing up to 30% of the water requirements of S. sempervirens each year (Burgess and Dawson 2004). For inclusion in this study, each site was required to have been previously clear-cut, be large enough for adequate sampling without edge effects (Russell and Jones 2001), and have not received post-harvest management such as seed- ing, thinning, or planting. Using these criteria, three study sites were selected in each of the five post-harvest age-classes (0-20, 21—40, 41—60, 81— 100, and 101-130 yr) as well as the three unharvested old-growth reference sites. The post-harvest age-class 61—80 yr was not sampled due to a lack of sites in that age range that met the criteria of this study. Sites were selected using detailed timber harvest and land management history maps on a GIS platform (Rutland 2002). Old-growth reference sites included Montgomery Woods State Natural Reserve (462 ha) located in the Big River watershed; Russell Unit (49 ha) of Mendocino Headlands State Park located in the Brewery Creek watershed adjacent to the Big River Unit; and Hendy Woods State Park (342 ha) located in the Navarro River watershed to the south of the Big River. These three sites were selected because they represent the only sizable remaining old-growth stands in Mendo- cino County. Data Collection Twenty, 0.031 ha (20 m diam.), circular sample | plots were randomly selected within each of the | 18 study sites, and located using a handheld GPS | receiver. Each sample plot was placed a minimum of 20 m from adjacent plots, 10 m from special | habitats such as riparian areas and rock outcrop- | pings, and 200 m from adjacent age-class | boundaries and main access roads. Plot size and © sampling intensity were determined through a | pilot study using the species-area curve method (Cain 1938) and are consistent with previous 2010] Pacific Ocean 0 1.25 29 iG. 1. 5 Miles research conducted in this forest type (Russell and Jones 2001; Loya and Jules 2007). Data collected on each plot included: tree canopy cover (measured at plot center using a spherical crown densiometer with one reading taken in each of the four cardinal directions); the occurrence and abundance of each tree species: the diameter (measured at 1.4 m above ground level) of all individuals greater than one meter in height; and the occurrence and abundance of all tree seedlings. The percent cover for all under- story species, including both herbs and shrubs, were determined using ocular estimates over the entire plot. In order to improve the accuracy of estimates plots were divided into eight sample wedges. Cover of each species was estimated in the field by two researches and averaged. Species were identified using the Jepson manual of higher plants of California (Hickman 1993). RUSSELL AND MICHELS: SEQUOIA SEMPERVIRENS CHRONOSEQUENCE Zo Legend pak hl Regenerating Sites Z | Big River Unit «vy, Russell redwood ” property * Montgomery Woods State Reserve se Hendy Woods State Park Location of sampled regenerating stands and old-growth reference sites in Mendocino County, California. Data Analysis As a preparatory procedure prior to conduct- ing ANOVA, we constructed a correlation matrix to examine possible relationships between stand characteristics and stand age. Significant correla- tions were found for several variables including tree density, seedling density, basal area and dominance of tree species, canopy cover, shrub cover, herbaceous cover, species richness, Shan- non-diversity (Weaver and Shannon 1949), and the cover of individual understory species includ- ing non-natives. One-way ANOVA analysis was used to test for differences among the means for each variable between age-classes and old-growth reference sites in a manner consistent with analysis of chronosequence data in other forest types (Pare and Bergeron 1995; Claus and George 2005; Delzon and Loustau 2005). Data 232 MADRONO [Vol. 57 TABLE 1. STAND CHARACTERISTICS OF SIX AGE-CLASSES IN A CHRONOSEQUENCE OF POST-HARVEST DEVELOPMENT IN THE CENTRAL RANGE OF THE SEQUOIJA SEMPERVIRENS FOREST. Age-classes sharing the same lower-case letter in each series were not significantly different, based on single factor ANOVA analysis (« = 0.05). Combined Redwood Tree density seedling density seedling density Basal area Richness Shannon- Age class (trees/ha) (seedlings/ha) (seedlings/ha) (m?/ha) (species/plot) diversity 0-20 yr 2048 a 890 a 184 a 14.8 a 21.9 a 2.0 a 21-40 yr 1889 a 11526 291a 22.9 b 18.4 b 2.0 a 41-60 yr 940 b 1243 b 3lla S36 18.6 b 22-0 81—100 yr 1260 c 1189 b 338 a 96.9 d 17.9 b 1EOve 101-130 yr 906 b 829 a 516 b 102.5 e 18.0 b 19c Old-growth 763 b 917 a,b 643 b 302.2 1 16.9 b hoiC were tested for homogeneity using the Bartlett’s Chi-Square statistic, and post-hoc analyses were conducted using the Bonferroni test for pair-wise differences between groups. Principle compo- nents analysis (PCA) was used to characterize general trends in species cover between age- classes. PCA data was transformed with individ- ual variable ranking to eliminate null values. Data analyses were conducted using Aable 2 statistical software (Gigawiz Ltd. Co., Tusla). RESULTS Tree Density, Dominance, and Diameter Distribution The density of trees (>1 m in height) declined with stand age (Table 1). The highest density was measured in the two youngest age-classes with significantly lower densities found in all other age-classes. Initial statistical equivalence with old-growth reference sites was reached in the third age-class suggesting the occurrence of a natural thinning event up until 40 yr. Somewhat higher densities on the two oldest age-classes, compared to the old-growth, suggest that stand thinning may continue at a reduced rate as the forest transitions toward old-growth conditions. The combined density of tree seedlings exhib- ited little variation between age-classes. A some- what higher number were found on sites ranging from 21-100 yr, however, all age-classes were statistically equivalent to old-growth. The density of S. sempervirens seedlings, however, exhibited a positive trend with stand age with the highest density found on the old-growth reference sites; statistical equivalence with old-growth was found for sites over 100 yr. No statistically significant relationship was found between stand age and the density of any other trees seedlings. The average combined basal area per hectare increased with stand age exhibiting significantly higher values in each successive age-class (Ta- ble 1). Basal area did not reach statistical equivalence with old-growth stands within the timeframe of the chronosequence. Analysis of the relative dominance (specific basal area/total basal area) of the three most common tree species (S-. sempervirens, L. densiflorus, and P. menziesii) indicated an increase in the relative dominance of S. sempervirens over time (Fig. 2), with S. sempervirens eclipsing all other species in the | | : 100% © © 80% © o 60% P, menziesii 8 ML. densiflorus S Ba MS. sempervirens | 5 x 20% - 0% 0-20 21-40 41-60 81-100 101-130 OG age-class FIG. 2. Relative dominance (specific basal area/total basal area) of Sequoia sempervirens, Lithocarpus densiflorus, — and Pseudotsuga menziesii across a 127-yr chronosequence of naturally regenerating Sequoia sempervirens stands. 2010] RUSSELL AND MICHELS: SEQUOIA SEMPERVIRENS CHRONOSEQUENCE 233 10000 >5-24cm 1000 1 25-49cm @ 50-99cm © ™ 100-149cm = 100 E @ 150-199cm 2 @ >200cm 10 1 0-20 41-60 81-100 101-130 OG age-class Fic. 3. Diameter distributions between six age-classes for all tree species combined on a _ post-harvest chronosequence in the central range of the Sequoia sempervirens forest. old-growth age class. The highest relative dom- inance for L. densiflorus was found in the two youngest age-classes, with significantly lower values found for all subsequent age-classes. The relative dominance of P. menziesii varied throughout the chronosequence with its highest values found in the 81—100 yr and 100-130 yr age-classes. An analysis of diameter distributions, based on size Classes defined by Guisti (2007), indicated an increase in the density of larger diameter trees over time (Fig. 3). The density of the smallest size-class of tree declined significantly in the early age classes, but showed no significant change after 40 yr. This result is consistent with a natural thinning event occurring early in the stand development process. Individual analysis of the diameter distributions of the three most common tree species (S. sempervirens, L. densiflorus, and P. menziesii) suggests a pulse of regeneration for each species early in the chronosequence (Fig. 4). It was also noted that S. sempervirens had the stems/ha 0-20 21-40 41-60 81-100 101-130 age-class 25-49cm stems/ha 21-40 41-60 81-100 101-130 age-class 50-99cm stems/ha 101-130 21-40 41-60 81-100 age-class stems/ha stems/ha 100-149cm ——— en 41-60 81-100 age-class en — ae 0-20 21-40 100+ OG 150-199cm a a 41-60 81-100 101-130 OG age-class >200cm stems/ha 41-60 81-100 age-class 101-130 Fic. 4. Diameter distribution of three species on six age-classes combined on a post-harvest chronosequence in the central range of the coast redwood forest (solid line = Sequoia sempervirens; wide-dashed line = densiflorus; narrow-dashed line = Pseudotsuga menziesii). Lithocarpus 234 highest density of trees in all size-classes across the chronosequence with the exception of the smallest size-class (O—24 centimeter) where L. densiflorus, and P. menziesii dominated, and the 25—49 centimeter) size class where P. menziesii dominated in the old-growth. Percent Cover of Canopy Layers The percent cover of trees, shrubs, and herbs varied significantly between age-classes (Fig. 5). The lowest tree cover was recorded in the two youngest age-classes with cover statistically equiv- alent to old-growth found in stands older than 81 yr. Shrub cover was highest in the two youngest age-classes, significantly lower in the 41-60 yr age-class, and progressively higher thereafter, reaching statistical equivalence with old-growth in the two oldest age-classes. Herb cover was uniformly low, and statistically equiv- alent, in all second-growth sites compared to old- growth where it was more than three times greater. Diversity and Species Distribution One hundred twenty-seven plant species were recorded in the sample plots (Appendix 1). The highest species richness (species/plot) was found in the youngest age-class with lower, statistically equivalent, values in all other age-classes includ- ing old-growth (Table 1). The Shannon-diversity index (H’) also exhibited a generally negative trend reaching statistical equivalence with old- growth in stands 81 yr and older. An H’ peak was found for the 41—60 yr age class, possibly in response to the natural thinning event noted in the earlier age-classes. Principal components analysis, using ranked percent cover of the 55 most common species, produced two axes that together explained 53.3% of the total variance with the first axis accounting for 35.9% and the second axis accounting for 17.4% (Fig. 6). The ordination illustrates group- ing by age-class along the x-axis (PC 1) with positive eigenvalues associated with the oldest age-classes including S. sempervirens (0.33), Trillium ovatum (0.45), Oxalis oregana (0.36), Viola sempervirens (0.34), Tiarella_trifoliata (0.32), and Calypso bulbosa (0.28); and negative eigenvalues associated with the younger age classes including Lithocarpus densiflorus (—0.31), Lonicera hispidula Douglas (—0.34), Whipplea modesta Torr. (—0.32), and Toxicodendron diver- silobum (Torr. & A. Gray) Greene (—0.27). The y-axis (PC_2) was positively associated with Vaccinium ovatum (0.41) and Rhododendron macrophyllum (0.30); and negatively correlated with Sequoia sempervirens (—0.31) and Oxalis oregana (—0.31). MADRONO [Vol. 57 Ten non-native plant species were recorded within the chronosequence (Table 2). The cover of each of these species declined with stand age to the extent that no non-native species were recorded in stands older than 60 yr. The absence of non- natives in the older age-classes suggests a lack of successful long-term establishment. However, the year of introduction, and historic distribution of each species, must also be considered as a possible explanation for their absence in older stands. While species such as Sonchus asper (L.) Hill and Stellaria media (L.) Vill. have been present in the region since the middle of the 1800’s, Cortaderia selloana (Schult.) Asch. & Graebn. and Leontodon leysseri (Vill.) M-rat may not have been present until the middle of the 1900's. Additional analysis of understory species cover indicated that several species commonly associ- ated with coast redwood forests increased with years since harvest. The cover of C. bulbosa, T. ovatum, and V. sempervirens increased to levels statistically equivalent to the old-growth refer- ence sites within the timeframe of the chronose- quence (Fig. 7a). The cover of Jris douglasiana, Tiarella trifoliata L., and Achlys triphylla also exhibited positive trends with stand age, but were significantly lower on all age-classes compared to old-growth (Fig. 7b). DISCUSSION In contrast to the “‘gap phase’’ succession process associated with old-growth coast red- wood forests, the second-growth stands studied in our post-harvest chronosequence followed patterns similar to forest types that regularly experience stand scale disturbance. Results indi- cate a progression through the four phases of succession outlined by Oliver (1981), “‘stand re- initiation,” “‘stem exclusion,” “‘understory re- initiation,” and “old-growth.” Many stand char- acteristics including tree density, canopy cover, shrub-cover, species diversity, non-native species occurrence, and the cover of several redwood associated species reached old-growth equiva- lence. While total basal area, dominance of S. sempervirens, herb cover, and the cover of several other redwood associated species, all progressed toward, but did not reach, old-growth equiva- lence within the 127-yr timeframe of the chrono- sequence. In addition, the diameter distribution of trees within age-classes indicated a transition from stands characterized by a high density of | small trees, to stands exhibiting a mixed size-class | distribution. These results are consistent with the | view that forests dominated by S. sempervirens have a high regenerative potential and are highly | resilient following disturbance (Allen et al. 1996). | The assertion that natural processes of com- | munity development are sufficient management approaches for the regeneration of coast redwood 2010] RUSSELL AND MICHELS: SEQUOIA SEMPERVIRENS CHRONOSEQUENCE 235 88 5 Tree 86 - 84 - 82 - 80 - 1S 76 - 74 12> 10.4 68 | % 0-20 21-40 41-60 81-100 101-130 OG % Shrub age-class % =X o1 ———_———— 0-20 21-40 41-60 81-100 101-130 OG 40 Herb age-class 0-20 21-40 41-60 ——— — a —y 81-100 101-130 OG age-class FIG. 5. Percent cover of trees, shrubs, and herbs on six age-classes on a post-harvest chronosequence in the central range of the Sequoia sempervirens forest; error bars indicate standard error. forests (Busing and Fujimori 2002, 2005) is also Supported by this research to some degree. The proliferation of Trillium ovatum, a species that is severely impacted by timber harvesting (Kahmen and Jules 2005), as well as several other coast redwood associates within the timeframe of our chronsequence, 1s encouraging. However, not all coast redwood-associated species recovered com- 236 MADRONO [Vol. 57 # 0-20 yrs + 21-40 yrs x 41-60 yrs oO 81-100 yrs O 101-130 yrs A Old-growth 300.00 200.00 100.00 she re) .00 a. -100.00 -200.00 -300.00 | 2 8 § 8 § 8 8 Bg 8 So r=) r=) So So So r=) So r=) ° ro) © =) ° ro) 5 N ae - N ise) rt Ww (To) PC_1 FIG. 6. Principle components for plant species across a 127-yr chronosequence of naturally regenerating Sequoia sempervirens stands. PC-1 explained 35.9% of variation. PC-2 explained an additional 17.4%. Convex outlines indicate post-harvest and old-growth age classes. pletely, suggesting that reestablishment of the coast redwood understory is a lengthy process. The ability to measure the full recovery of S. sempervirens stands following timber harvest was limited by the length of our chronosequence in relation to the life span of the dominant organism (>1500 yr). The study was also limited by gathering data exclusively at ground level. TABLE 2. Considering the volume and complexity of the old-growth coast redwood canopy, the full development of a stand may require several centuries (Sillett and Baily 2003). The study of post-harvest patterns of canopy development, as well as analysis of soil organism assemblages and wildlife habitat features, could increase insight into the long-term effects of timber harvest, and NON-NATIVE PLANT SPECIES RECORDED IN SIX AGE-CLASSES IN THE CENTRAL RANGE OF THE COAST REDWOOD FOREST. The estimated date of introduction is based on the earliest specimen records for each species in northern coastal California, retrieved from the Consortium of California Herbaria (http://ucjeps.berkeley.edu/ cgi-bin/get_consort). Species Arabidopsis thaliana (L.) Heynh. (Mouse Ear Cress) Cirsium vulgare (Savi) Ten. (Common Bull Thistle) Cortaderia selloana (Schult.) Asch. & Graebn. (Pampas Grass) Hypochaeris glabra L. (Smooth Cat’s Ear) Hypochaeris radicata L. (Hairy Cat’s Ear) Lactuca saligna L. (Willowleaf Lettuce) Leontodon taraxacoides (Vill.) Mérat (White-Flowered Hawk Bit) Sonchus asper (L.) Hill (Spiny Sow Thistle) Stellaria media (L.) Vill. (Common Chickweed) Taraxacum officinale F. H. Wigg. (Dandelion) Estimated date of Oldest age-class introduction present 1926 41-60 1900 41-60 1941 41-60 1888 41-60 1900 41-60 1927 0-20 1938 21-40 1861 21-40 1876 21-40 1895 0-20 2010] RUSSELL AND MICHELS: SEQUOIA SEMPERVIRENS CHRONOSEQUENCE 23) 1.4 - 125 1 Be ees “a --| Calypso bulbosa oO 0.8 - Ares Bn Trillium ovatum 5 oe A — —-—-Viola sempervirens 5 (0.6 - vs oS ee ee 0.4 - oe ee = 0.2 - ee 0-20 21-40 41-60 81-100 101-130 OG age-class 16 1.4 5 es a lris douglasiana © a oak Tiarella trifoliata Oe — — —-Achlys triphylla Ss 0.6 + 0.4 - 0.2 | fee cern Q) {a — 0-20 21-40 41-60 81-100 101-130 OG age-class FIG. 7. Mean percent cover of coast redwood associated species on six post-harvest age-classes on a post-harvest chronosequence in the central range of the Sequoia sempervirens forest; error bars indicate standard error; a) represents species that reached OG equivalence, b) represents species that did not reach OG equivalence. the regenerative potential of the entire forest community over time. ACKNOWLEDGMENTS This research was supported by a grant from the Save-the-Redwoods League, 114 Sansome Street, Suite 1200. San Francisco, California. Logistical support was provided by Rene Pasquinelli and Bill Maslach of the California Department of Parks and Recreation, Marc Jameson, Rob Rutland, and Lynn Webb of the California Department of Forestry and Fire Protection, and Cal Winslow of the Mendocino Institute. 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Don 0.00 Pinus sabiniana Douglas 0.00 Pseudotsuga menziesii (Mirb.) Franco var. menziesii 0.04 Salix scouleriana Hook. <0.01 Sequoia sempervirens (D. Don) Endl. 0.14 Taxus brevifolia Nutt. <0.01 Torreya californica Torr. <0.01 Tsuga heterophylla (Raf.) Sarg. <0.01 Umbellularia californica (Hook & Arn.) Nutt. 0.02 Shrubs (% cover/plot) Arctostaphylos columbiana Piper 1.08 Baccharis pilularis DC. 0.00 Berberis aquifolium Pursh 0.00 Age class 21—40 41—60 8 1—100 101-130 OG 0.00 0.04 0.26 <0.01 0.02 0.01 0.01] <0.01 0.04 0.00 <0.01 0.00 0.05 <0.01 0.00 0.00 0.00 0.00 0.00 <0.01 0.16 0.11 0.20 0.21 <0.01 = (0:01 <0.01 <0.01 0.00 <0.01 0.00 <0:0] 0.12 <0.01 0.01 0.00 0.00 0.00 =<0.01 0.00 0.02 O17 0.44 0.49 0.17 0.00 0.00 0.00 <0.01 0.00 0.50 1.26 2.06 2213 10.92 0.00 0.00 0.00 <0.01 0.00 <0.01 0.00 0.00 0.00 0.00 = 0,01 Oe es 0.03 0.09 0.07 0.01 <0.01 <0.01 0.02 0.05 1.30 0.00 0.00 0.00 0.00 1.30 0.00 0.00 0.00 0.00 0.00 0.00 0.08 0.00 0.03 240 MADRONO [Vol. 57 APPENDIX 1. CONTINUED. Age class 0-20 21-40 41—60 81-100 101-130 OG Berberis nervosa Pursh 0.12 0.57 0.50 0.11 0.24 0.10 Blechnum spicant (L.) Sm. 0.00 0.15 0.10 0.43 0.10 1.18 Ceanothus thyrsiflorus Eschsch. 6.82 1.02 O22 0.00 0.02 0.00 Euonymus occidentalis Torr. 0.00 0.00 0.00 0.00 0.00 0.09 Gaultheria shallon Pursh 0.18 0.54 0.15 2.48 1.62 Revd Lepechinia calycina (Benth.) Epling 0.00 0.00 0.00 0.00 0.01 0.00 Lonicera hispidula Douglas 0.66 0.48 0.66 0.38 0.15 O22 Rhododendron macrophyllum D. Don 1.24 1.24 0.38 2.46 1.78 4.28 Ribes menziesii Pursh 0.09 0.03 0.00 0.03 0.00 0.02 Rosa gymnocarpo Nutt. 0.05 0.08 0.13 0.03 0.05 0.20 Rubus leucodermis Torr. & A. Gray O32 0.23 0.22 0.00 0.00 0.35 Rubus parviflorus Nutt. 1.02 0.32 0.43 0.00 0.02 0.00 Rubus spectabilis Pursh 0.00 0.00 0.00 0.00 0.00 0.05 Rubus ursinus Cham. & Schldl. 0.24 0.14 1.00 0.14 0.01 0.04 Symphoricarpos albus (L.) S. F. Blake 0.00 0.00 0.01 0.00 0.02 0.00 Toxicodendron diversilobum (Torr. & A. Gray) Greene 5 es 1.04 0.80 0.47 0.58 [53 Vaccinium ovatum Pursh 6.82 5.3) LST 5167 4.71 |e Vaccinium parvifolium Sm. 0.38 Q.27 0.30 0.74 0.39 235 Ferns (% cover/plot) Adiantum aleuticum (Rupr.) C.A. Paris 0.07 0.05 0.16 0.10 0.26 0.02 Athyrium filix-femina (L.) Roth 0.00 0.05 0.16 0.10 0.26 0.02 Dryopteris arguta.(Kaulf.) Maxon O27 0.16 0.68 0.08 0.00 0.09 Pentagramma triangularis (Kaulf.) Yatsk., Windham & E. Wollenw. 0.58 0.18 0.10 O.02 0.02 0.02 Polypodium californicum Kaulf. 0.00 0.00 0.00 0.06 0.00 0.09 Polystichum munitum (Kaulf.) C. Presl 6.24 2.59 4.07 4.10 6.01 5.00 Pteridium aquilinum var. pubescens L. Underw. 0.93 1.14 0:55 1.06 0.39 1.62 Woodwardia fimbriata Sm. 0.00 0.00 0.00 0.03 0.00 3.08 Herbs (% cover/plot) Achlys triphylla (Sm.) DC. 0.03 0.18 0.10 0.00 0.35 0.87 Actaea rubra (Aiton) Willd. 0.02 0.01 0.09 0.02 0.15 0.11 Adenocaulon bicolor Hook. 0.05 0.00 0.08 0.08 0.07 0.80 Agoseris retrorsa (Benth.) Greene 0.01 0.05 0.00 0.00 0.00 0.00 Anaphalis margaritacea (L.) Benth. & Hook. 0.03 0.38 0.00 0.00 0.00 0.00 Anemone deltoidea Hook. 0.00 0.00 0.00 0.00 0.00 0.02 Aquilegia formosa Fisch. 0.03 0.00 0.02 0.00 0.00 0.22 Arabidopsis thaliana (L.) Heynh. 0.04 0.04 0.03 0.00 0.00 0.00 Aralia californica 8. Watson 0.00 0.00 0.00 0.05 0.00 0.08 Asarum caudatum Lindl. 0.12 0.27 0.05 0.03 0.63 0.36 Calypso bulbosa (L.) Oakes 0.02 0.03 0.17 0.16 0.22 0.09 Calystegia occidentalis (A. Gray) Brummitt 0.00 0.11 0.00 0.00 0.00 0.00 Campanula prenanthoides Durand 0.17 0.07 0.18 0.01 0.02 0.11 Cardamine californica var. californica (Torr. & A. Gray) Greene 0.06 0.07 0.18 0.76 O25 0.27 Chenopodium berlandieri Moq. 0.02 0.00 0.00 0.00 0.00 0.00 Chimaphila menziesii (D. Don) Spreng. 0.00 0.00 0.01 0.10 0.05 0.01 Chlorogalum pomeridianum (DC.) Kunth 0.01 0.00 0.07 0.00 0.00 0.00 Cirsium vulgare (Savi) Ten. 0.13 0.06 0.18 0.00 0.00 0.00 Claytonia perfoliata Willd. 0.00 0.02 0.04 0.00 0.00 0.00 Claytonia sibirica L. 0.01 0.00 0.01 0.01 0.07 0.00 Clintonia andrewsiana Torr. 0.05 0.05 0.03 0.35 0.07 O72 Collomia heterophylla Hook. 0.00 0.01 0.00 0.00 0.00 0.00 Corallorhiza maculata Rat. 0.00 0.00 0.02 0.16 0.06 0.03 Corallorhiza striata Lindl. 0.00 0.00 0.01 0.01 0.02 0.00 Cordylanthus tenuis A. Gray spp. tenuis 0.01 0.00 0.00 0.00 0.00 0.00 Cortaderia selloana (Schult.) Asch. & Graebn. 0.10 0.28 0.13 0.00 0.00 0.00 Cynoglossum grande Lehm. 0.02 0.02 0.03 0.10 0.00 0.00 2010] RUSSELL AND MICHELS: SEQUOIA SEMPERVIRENS CHRONOSEQUENCE 24] APPENDIX |. CONTINUED. Age class 0-20 21—40 41-60 81—100 101-130 OG Dicentra formosa (Haw.) Walp. 0.17 0.13 0.01 0.00 0.18 0.00 Disporum hookeri (TYorr.) G. Nicholson 0.37 0.25 0.49 0.39 0.35 1.53 Epilobium angustifolium L. 0.03 0.13 0.84 0.00 0.00 0.00 Epilobium ciliatum Raf. 0.01 0.00 0.03 0.00 0.00 0.00 Equisetum arvense L. 0.04 0.00 0.02 0.03 0.07 0.05 Fragaria chiloensis (L.) Duchesne 0.05 0.00 0.03 0.00 0.00 0.00 Galium californicum Hook. & Arn. 0.03 0.07 0.00 0.00 0.00 0.22 Galium triflorum Michx. 0.36 0.23 0.61 0.41 0.23 0.72 Goodyera oblongifolia Raf. 0.00 0.00 0.00 0.20 0.00 0.02 Hemizonia corymbosa (DC.) Torr. & A. Gray 0.00 0.00 0.04 0.00 0.00 0.00 Holodiscus discolor (Pursh) Maxim. 0.03 0.00 0.00 0.00 0.00 0.00 Hypochaeris glabra L. 0.03 0.03 0.01 0.00 0.00 0.00 Hypochaeris radicata L. 0.00 0.00 0.02 0.00 0.00 0.00 Tris douglasiana Herb. 0.00 0.02 0.02 0.04 0.03 0.13 Lactuca saligna L. 0.01 0.00 0.00 0.00 0.00 0.00 Lathyrus vestitus Nutt. 0.05 0.00 0.02 0.00 0.00 0.03 Leontodon taraxacoides (Vill.) Mérat 0.01 0.03 0.00 0.00 0.00 0.00 Lithophragma glabrum Nutt. 0.00 0.00 0.00 0.00 0.00 0.00 Lotus purshianus Clem. & E.G. Clem. 0.00 0.00 0.00 0.00 0.01 0.00 Lotus stipularis (Benth.) Greene 0.32 0.14 0.22 0.04 0.03 0.04 Mimulus aurantiacus Curtis 0.12 0.00 0.00 0.00 0.00 0.00 Nemophila menziesii Hook. & Arn. 0.01 0.00 0.00 0.00 0.00 0.00 Nemophila parviflora Benth. 0.05 0.00 0.02 0.00 0.00 0.00 Oxalis oregana Nutt. 1.68 0.97 0.77 1.63 3.00 23,27 Petasites frigidus (L.) Fries 0.02 0.00 0.00 0.00 0.00 0.00 Polygala californica Nutt. 0.11 0.00 0.01 0.02 0.00 0.01 Prunella vulgaris L. 0.00 0.00 0.00 0.00 0.00 0.00 Pyrola picta Sm. 0.03 0.00 0.00 0.00 0.00 0.00 Rhamnus alnifolia L’ Her. 0.01 0.05 0.00 0.04 0.00 0.02 Rhamnus purshiana DC. 0.00 0.00 0.00 0.00 0.00 0.02 Sanicula crassicaulis DC. 0.02 0.00 0.02 0.00 0.00 0.00 Satureja douglasii (Benth.) Briq. 0.05 0.03 0.18 0.00 0.00 0.00 Scoliopus bigelovii Torr. 0.00 0.00 0.00 0.03 0.00 0.00 Scrophularia californica Cham. & Schldl. 0.02 0.20 0.00 0.00 0.00 0.00 Senecio vugares L. 0.00 0.02 0.00 0.00 0.00 0.00 Smilacina racemosa (L.) Link 0.11 0.00 0.04 0.05 0.04 0 Smilacina stellata (L.) Desf. 0.05 0.04 0.05 0.00 0.04 0.26 Sonchus asper (L.) Hill 0.03 0.04 0.00 0.00 0.00 0.00 Stachys bullata Benth. O27 0.23 0.24 0.00 O02 0.03 Stachys ajugoides Benth. 0.00 0.00 0.00 0.08 0.00 0.58 Stellaria crispa Cham. & Schldl. 0.01 0.00 0.00 0.00 0.00 0.00 Stellaria media (L.) Vill. 0.00 0.01 0.00 0.00 0.00 0.00 Stephanomeria exigua Nutt. 0.00 0.00 0.00 0.00 0.00 0.00 Taraxacum officinale F. H. Wigg. 0.02 0.00 0.00 0.00 0.00 0.00 Tiarella trifoliata L. 0.01 0.00 0.06 0.10 0.16 1.84 Trientalis latifolia Hook. 0.83 0.42 0.20 0.63 0.23 0.42 Trillium chloropetalum (Torr.) Howell 0.04 0.00 0.00 0.05 0.00 0.50 Trillium ovatum Pursh 0.28 0.38 0.36 0.86 0.52 1:07 Urtica dioica subsp. holosericea (Nutt.) Thorne 0.00 0.00 0.00 0.00 0.02 0.00 Vancouveria planipetala Calloni 0.46 0.29 0.21 O33 0.18 0.01 Veratrum californicum Durand 0.00 0.00 0.00 0.03 0.00 O22 Vicia americana Muhl. ex Willd. 0.01 0.00 0.00 0.00 0.00 0.00 Viola glabella Nutt. 0.00 0.00 0.00 0.02 0.08 0.93 Viola sempervirens Greene 0.28 0.40 0.53 0.96 0.44 1.03 Whipplea modesta Torr. LAS 1.15 1.28 0.20 O23 0.19 Yabea microcarpa (Hook. & Arn.) Koso-Pol. 0.03 0.03 0.00 0.00 0.00 0.00 MADRONO, Vol. 57, No. 4, pp. 242-245, 2010 REDISCOVERY OF PLAGIOBOTHRYS HYSTRICULUS (BORAGINACEAE) AND NOTES ON ITS HABITAT AND ASSOCIATES ROBERT E. PRESTON, BRAD D. SCHAFER, AND MARGARET WIDDOWSON ICF International, 630 K Street, Suite 400, Sacramento, CA 95814 rpreston@icfi.com ABSTRACT The rediscovery of bearded popcorn flower, the last confirmed collection of which was in 1892, is documented. The species is endemic to Napa, Solano, and Yolo counties, California, where it grows in vernal pools, vernal swales, and other moist areas in grasslands. Associated species include wetland and upland grasses and forbs typically found in the vernal pool-grassland ecotone. Key Words: Boraginaceae, endemism, Plagiobothrys hystriculus, rare species, vernal pools. Bearded popcorn flower, Plagiobothrys hystri- culus (Piper) I. M. Johnst., is a rare plant previously confirmed from only two collections from Solano Co. The last known confirmed collection was made in 1892 by Willis Jepson, whose specimens became the type of A/locarya hystricula Piper (Piper 1920). Jepson’s specimens cited the Montezuma Hills as the general collection location but provided no other detail. The other collection was made by Katherine Brandegee in 1883. Brandegee cited her collection locality as Elmira, north of the present Jepson Prairie, where she collected many grassland and vernal pool plants. The exact location of her collection site is unknown; all of the land in the vicinity of Elmira has long since been converted to agriculture. One other partial specimen apparently collected by K. Brandegee exists (UCI101495), although no date or location information 1s associated with the specimen. Because Plagiobothrys hystriculus was known only from two historic collections and no subse- quent confirmed observations of it had been made, the California Native Plant Society (2001) had listed the species as extinct. Numerous attempts by botanists over many years (including the authors of this report) to relocate the species in the Montezuma Hills area apparently had been unsuccessful (T. Messick, ICF International, personal communication). Nevertheless, P. Aystri- culus was one of the 33 plant and animal species associated with vernal pools that were included in the ““Recovery Plan for Vernal Pool Ecosystems of California and Southern Oregon” (United States Fish and Wildlife Service 2005). Although not listed as threatened or endangered, it was included in the Recovery Plan to put conservation actions in place should it be rediscovered. REDISCOVERY In May 2005, B. Schafer and M. Widdowson rediscovered and collected Plagiobothrys hystri- culus in the Montezuma Hills during pre-project botanical surveys. The specimen was seen and identification confirmed by T. Messick, author of the Plagiobothrys treatment for the Jepson Manual (Messick 1993); the specimen was also compared with the isotype housed at the Jepson Herbarium, making this the first confirmed observation in almost 114 yr. Based on Schafer and Widdowson’s description of the habitat in which the plants were found, R. Preston located two additional populations during surveys for an adjacent project. In 2006, after confirming that the species was present at the Montezuma Hills localities, Preston conducted a search for Plagi- Obothrys hystriculus at two vernal pool preserves (Wilcox Ranch, Jepson Prairie) located geo- graphically intermediate between the Montezuma Hills and Elmira, successfully finding populations at both sites. Although ours are the first confirmed collec- tions of Plagiobothrys hystriculus since 1892, we are not the first to recollect the species. A subsequent search of the herbarium at the University of California, Davis (DAV) and the © University of California/Jepson Herbaria (UC/ | JEPS) turned up additional specimens that had © been misdetermined as P. acanthocarpus (Piper) I. M. Johnst., P. greenei (A. Gray) I. M. Johnst. or | P. glyptocarpus (Piper) I. M. Johnst. The late Professors Beecher Crampton and John Tucker, | both botanists at U. C. Davis, collected P. hystriculus specimens at or near Jepson Prairie in the 1960’s and 1970’s. Bob Holland, the noted | vernal pool authority, also collected P. hystriculus | at Jepson Prairie in 1976 and again in 1981. Jake | Ruygt collected specimens in Napa Co. in 1998 | while collecting for his Napa County flora project, and Ayzik Solomeshch collected P. hystriculus 1. 2002 while sampling vernal pools | in Solano Co. Why has this species been so difficult to locate? © There are several likely explanations. Most of the | land in the Montezuma Hills is privately owned | 2010] and not accessible for searches. The survey window for the species appears to be very narrow, as the blooming period lasts for only about three weeks, from the last week of April to the second week of May, and the plants are well into fruit and senescing by mid-May. The plants themselves are low and spreading, sometimes growing under a dense cover of Lolium multiflorum Lam., making detection extremely difficult. The plants are locally uncommon, occurring in small scattered stands, typically consisting of 10 to 20 plants. Annual variation in rainfall may also affect the ability of field workers to locate this species during any given year. The higher than average rainfall in 2005 and 2006 may have created optimal conditions for observing the species. DISTRIBUTION As currently known, the primary range of Plagiobothrys hystriculus consists of an approxi- mately 150 square-mile area in central Solano Co., bordered to the south by the Montezuma Hills, to the north by Alamo and Ulatis creeks, and to the east by the Yolo Bypass. Another disjunct population occurs in Napa Co. The elevation range is from about 5 ft above sea level to about 870 ft above sea level. Although these observations have considerably expanded the known distribution of P. Aystriculus, the species remains extremely rare and should remain a species of conservation concern. Fortunately, many of the new occurrences are located in vernal pool preserves. HABITAT AND ASSOCIATES Previous habitat characterizations described the habitat of Plagiobothrys hystriculus as “low plains” (Jepson 1925) or ‘‘grassy hillsides and plains” (Abrams 1951). Messick (1993) indicated that the habitat probably consisted of vernal pools or other wet sites, similar to other species in Plagiobothrys section Allocarya. Our observa- tions indicate that P. Aystriculus occurs in vernal pools and vernal swales and also in other vernally moist areas in grasslands that do not pond for significant duration but have saturated soil for long periods during the rainy season. Plants associated with P. hystriculus are those species commonly found at the ecotone between vernal pools and the adjacent annual grassland, includ- ing both native and introduced grasses and forbs (Table 1). Most documented populations of Plagio- bothrys hystriculus occur where soils have been mapped as San Ysidro sandy loam or Solano loam (Bates 1977). These soil series have increased clay content in the subsoil, which results in very slow permeability, and wetlands occur where swales or basins are present. One PRESTON ET AL.: REDISCOVERY OF PLAGIOBOTHRYS HYSTRICULUS 243 population is on soil mapped as Rincon clay loam and another is on soil mapped as Capay clay (Bates 1977). The latter soil series have slow permeability but are less likely to support wetlands. MORPHOLOGICAL COMPARISONS The Jepson Manual key to the species of Plagiobothrys (Messick 1993) generally works well for section Allocarya. However, persons unfamiliar with the nutlet morphology or with the corresponding terminology could experience difficulty using the key. Plagiobothrys hystriculus is one of only four species (P. acanthocarpus, P. austinae [Greene] I. M. Johnst., P. greenei, P. hystriculus) that have prickles, large (ca. | mm) spine-like projections, on the abaxial nutlet surface (Fig. lb-d). The prickles on these species are also beset with short, hair-like bristles that spread at right angles or curve towards the base of the prickles. Several other Plagiobothrys species have nutlets with bristles . (P. leptocladus [Greene] I. M. Johnst., P. hispidulus [Greene] I. M. Johnst., P. humistratus I. M. Johnst., P. scriptus [Greene] I. M. Johnst.); however, all of these species lack prickles. Plagiobothrys glyptocarpus (Fig. la) and P. trachycarpus 1. M. Johnst. (Fig. le) sometimes have very short prickles (ca. 0.1— 0.2 mm) lacking bristles. KEY TO THE PLAGIOBOTHRYS SPECIES WITH BOTH PRICKLES AND BRISTLES 1. Prickles on nutlet margins and abaxial ridge only; nutlet surface smooth, ridges or papillae ODSCINe = a ee eee P. austinae 1’ Prickles evenly distributed abaxially on nutlet; nutlet surface with ridges and/or papillae 2. Nutlet surface prominently ridged abaxially and adaxially, the spaces between ridges with coarse papillae ....... P. acanthocarpus 2’ Nutlet surface lacking ridges or with obscure adaxial ridges, finely papillate 3. Nutlet surface papillae glabrous, ridges absent; bristles arched towards base of prickles; only lowermost flowers of inflorescence bracted.. ......%. P. greenei Nutlet surface papillate, papillae dense- ly bristled, adaxial surface with obscure ridges; bristles short, straight; inflores- cence bracted throughout... . P. hystriculus ~ 3 SPECIMENS EXAMINED The following collections summarize the known localities for Plagiobothrys hystriculus (additional collections/duplicates not listed have been distributed). USA. CALIFORNIA. Napa Co.: 4.7 mi N of first bridge on Berryessa- Knoxville Rd, J. Ruygt 3981] (JEPS). Solano Co.: Montezuma Hills, W. L. Jepson 21176 (JEPS, 244 TABLE 1. SOLANO Co., CALIFORNIA. Species Lolium multiflorum Lam. Lythrum hyssopifolium L. Juncus bufonius L. Hordeum marinum Hudson subsp. gussoneanum (Parl.) Thell. Achyrachaena mollis Schauer Eryngium aristulatum Jepson Crassula aquatica (L.) Schonl. Plagiobothrys stipitatus (Greene) I. M. Johnst. Geranium dissectum L. Bromus hordeaceus L. Cicendia quadrangularis (Lam.) Griseb. Plagiobothrys greenei (A. Gray) I. M. Johnst. Psilocarphus tenellus Nutt. Rumex crispus L. Briza minor L. Lasthenia fremontii (A. Gray) Greene Plagiobothrys bracteatus (TY. J. Howell) I. M. Johnst. Plantago coronopus L. Pogogyne zizyphoroides Benth. Ranunculus muricatus L. Trifolium depauperatum Desvy. Triphysaria eriantha (Benth.) T. I. Chuang & Heckard Cotula coronopifolia L. Deschampsia danthonioides (Trin.) Munro Downingia concolor Greene Erodium botrys (Cav.) Bertol. Limnanthes douglasii R. Br. Phalaris lemmonii Vasey Trifolium dubium Sibth. Veronica peregrina L. Aira caryophyllea L. Avena barbata Link Brodiaea elegans Hoover Convolvulus arvensis L. Holocarpha virgata (A. Gray) Keck Lasthenia californica Lindl. Lasthenia glaberrima A DC. Lotus corniculatus L. Plagiobothrys leptocladus (Greene) I. M. Johnst. Pleuropogon californicus (Nees) Benth. ex Vasey Poa annua L. Psilocarphus oregonus Nutt. Trifolium gracilentum Torr. & A. Gray Vulpia myuros (L.) C. C. Gmel. isotype); Elmira, K. Brandegee, May 1883 (UC); 2 mi NE of Dozier Station, at jct Brown and Robbens roads, B. Crampton 6334 (AHUC); 2 mi SW of Dozier Station, B. Crampton 8700 (UC, AHUC); on N side of small road running W from intersection of Cook Lane and Sacramento Northern Railroad, ca. 1/2 mi W of. this intersection, J. M. Tucker 4348 (DAV); N of Alkali Lake, Dozier, R. Holland 167 (UC); Jepson Prairie TNC Preserve, Section 23, R. Holland 1082 (UC); Gridley Ranch, 1 mi N from intersection of Hastings Rd and Salem Rd, A. Solomeshch, 27 Apr 2002 (DAV); Montezuma Hills, approximately | mi S of the junction of CA MADRONO [Vol. 57 SPECIES OCCURRING WITH PLAGIOBOTHRYS HYSTRICULUS, RECORDED IN 24 1-M DIAMETER PLOTS IN Frequency 95.83% 83.33% 62.50% 50.00% 41.67% 37.50% Pied Whe 25.00% 20.83% 16.67% 16.67% 16.67% 16.67% 16.67% 12.50% 12.50% 12.50% 12.50% 12.50% 12.50% 12.50% 12.50% 8.33% 8.33% §.337% 8.33% 8.33% 8.33% 8.33% 8.33% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% 4.17% Hwy 12 and CA Hwy 113, Brad D. Schafer 135 & | Margaret Widdowson (JEPS), Brad D. Schafer | 137 & Margaret Widdowson (DAV); Montezuma | Hills, approximately 1.5 mi S of Hwy 12 and E of | Olsen Rd, R. E. Preston 2347 (JEPS); Kirby Hill, | at toe of north slope, R. E. Preston 2348 (DAY); 4.6 mi E of Travis Air Force Base, at Wilcox | Ranch Preserve, R. E. Preston 2383 (DAV); 3 mi | E of Travis Air Force Base, at Wilcox Ranch | Preserve, R. E. Preston 2389 (DAV); Jepson | Prairie Preserve, 1.4 mi SSW of the intersection of | Cook Lane and Hwy 113, R. E. Preston 2393 (DAV). Yolo Co.: DFG Tule Ranch, C. W. Witham 1562 (DAVY). 2010] PRESTON ET AL.: REDISCOVERY OF PLAGIOBOTHRYS HYSTRICULUS 245 Fic. 1. greenei. d. P. hystriculus. e. P. trachycarpus. Scale bars = ACKNOWLEDGMENTS We thank Tim Messick for determination of the specimens from the initial rediscovery site; Solano Land Trust for access to Wilcox Ranch and Jepson Prairie Preserves; Rocky Turcotte and Laurie Preston for field assistance; Jake Ruygt and Carol Witham for providing unaccessioned specimens; and the staff of the UC/JEPS and AHUC/DAV herbaria for access to their speci- mens. LITERATURE CITED ABRAMS, L. 1951. Illustrated flora of the Pacific States, Volume III: Geraniaceae to Scrophulariaceae. Stanford University Press, Stanford, CA. BATES, L. A. 1977. Soil survey of Solano County, California. U.S. Department of Agriculture, Soil Conservation Service, in cooperation with Univer- sity of California Agricultural Experiment Station, Washington, DC. Comparison of the nutlets of five Plagiobothrys species. a. P. glyptocarpus. b. P. acanthocarpus. c. P. 3.0 mm. CALIFORNIA NATIVE PLANT SOCIETY. 2001, Inventory of rare and endangered plants of California (sixth edition). Rare Plant Scientific Advisory Commit- tee, David P. Tibor, Convening Editor. California Native Plant Society, Sacramento, CA. JEPSON, W. L. 1925. A manual of the flowering plants of California. Associated Students Store, Berkeley, CA. Messick, T. C. 1993. Plagiobothrys. Pp. 386—390 in J.C. Hickman (ed.), The Jepson manual: higher plants of California. University of California Press, Berkeley, CA. PIPER, C. V. 1920. A study of A//ocarya. Contributions from the U.S. National Herbarium 22:79-113. UNITED STATES FISH AND WILDLIFE SERVICE. 2005, Recovery plan for vernal pool ecosystems of California and southern Oregon. U.S. Fish and Wildlife Service, Portland, OR. Website http://www. fws.gov/sacramento/es/recovery_plans/vp_recovery_ plan_links.htm [accessed 28 December 2010]. MADRONO, Vol. 57, No. 4, pp. 246-260, 2010 TAXONOMIC NOVELTIES FROM WESTERN NORTH AMERICA IN MENTZELIA SECTION BARTONIA (LOASACEAE) JOHN J. SCHENK' AND LARRY HUFFORD School of Biological Sciences, P.O. Box 644236, Washington State University, Pullman, WA 99164-4236 jschenk@bio.fsu.edu ABSTRACT Recent field collections and surveys of herbarium specimens have raised concerns about species circumscriptions and recovered several morphologically distinct populations in Mentzelia section Bartonia (Loasaceae). From the Colorado Plateau, we name M. paradoxensis from Paradox and Gypsum valleys of western Colorado, which is closely related to M. marginata. We name M. holmgreniorum from northeastern Arizona and M. filifolia from the northern border region of Arizona and New Mexico, both of which share morphological similarities with M. /agarosa, M. laciniata, and M. conspicua. From north central New Mexico, we name M. sivinskii, which is most closely related to M. procera and M. integra. We describe three varieties of M. /ongiloba, including M. longiloba var. yavapaiensis, which is distributed throughout Arizona, M. longiloba var. pinacatensis, which is narrowly distributed in the Pinacate Desert of Sonora, Mexico, and the northern Chihuahuan Desert M. longiloba var. chihuahuaensis. We propose the new combinations M. lagarosa and M. procera to alleviate the polyphyly of M. pumila. Key Words: Cryptic species, intermountain West, new species, polyphyletic taxa, systematics. Mentzelia section Bartonia (Loasaceae) is a monophyletic group (Hufford 2003; Hufford et al. 2003; Schenk 2009) dispersed throughout the arid North American West. The taxonomy of the section and collection limitations (Thompson and Prigge 1986) have long encumbered a clear understanding of the biological diversity of the group. Collections made in the last two decades have expanded our knowledge of diversity in the section and resulted in the description of 15 new species. Many of these recently described species are associated with restrictive substrates (e.g., gypsum-rich soils) and are narrowly distributed (Prigge 1986; Holmgren and Holmgren 2002; Schenk et al. 2010). As part of a revisionary study of Mentzelia section Bartonia, we have made extensive new collections of the group, especially to examine the distribution and circumscriptions of poorly de- limited taxa. This fieldwork and an accompany- ing survey of herbarium collections indicated that some populations possess morphological states that could not be readily accommodated in existing taxa in section Bartonia. The possibility that the existing taxonomy of the section does not capture the diversity that is present among natural populations has led us to sample distinc- tive populations as part of phylogeny reconstruc- tions based on molecular data to examine their relationships to better known species (Schenk 2009). We have used insights from the fieldwork, ‘Current address: Department of Biological Science, Florida State University, Tallahassee, FL 32306-4295. survey of herbarium specimens, and the phyloge- netic placements of distinctive populations as a guide to their taxonomic treatment. Our results indicate that several populations are independent evolutionary lineages that do not readily fit existing species circumscriptions, and we describe these entities as new species. Several evolutionary lineages found to be associated with M. longiloba J. Darl. are morphologically and geographically distinctive (Schenk 2009), and we describe these as varieties of M. longiloba. In addition, our related | molecular phylogenetic studies (Schenk 2009) also © indicate that M. pumila Torr. & A. Gray as treated by Darlington (1934) and Thorne (1986) is | polyphyletic. In order to alleviate the polyphyly of | M. pumila, we make two new combinations. MATERIALS AND METHODS Field observations of nearly all species of Mentzelia section Bartonia and a study of our | recent collections and specimens, including types, » in herbaria were used to assess population and | species variation. Morphological measurements were taken with digital calipers, using a dissecting | microscope when necessary. Leaf characters were | measured separately for leaves on the lower 1/3 © and upper 1/3 of main stem or renewal axes. Leaf measurements were not taken from secondary or — higher order (lateral, nonrenewal) branches | because they often differed in size and shape | from those on main stems. Leaf lengths were | measured from the distal tip of the lamina to the | junction of the leaf base with stem. Leaf widths were measured at the widest point of the lamina. | 2010] The rachis width was measured as the distance across the sinuses at the shortest length between lobes (if present) and at the widest point of the lamina. Lobe width (when lobes were present), a measure across lobes, was measured at mid- length for a lobe positioned at the widest point of the leaf. The number of lobes per leaf included the total number of lobes on both margins of the leaf. Trichome densities and composition on leaves were characterized for both abaxial and adaxial surfaces of the lamina, excluding the central vein and margins. Prophylls_ either subtended all flowers or were adnate to the ovary. Prophyll measurements were made for the most distal bract that either subtended a flower or was adnate to a flower ovary. Calyx lobe lengths were measured from the base of a lobe to its distal tip, excluding the calyx tube and hypanthium. Petal lengths were measured from the base of a petal to its distal tip. Petal and median antesepalous (=outermost) stamen widths were measured at their widest points. We characterized petals as narrowly to broadly spatulate, oblanceolate, or elliptic based on assessment of overall shape. Elliptical (broadest near mid-length) and oblanceolate (broadest distal to mid-length) petals largely lacked a differentiated claw and limb in contrast to spatulate petals, which had distinct claw and limb regions. Styles were measured from the top of the ovary to the tip of the stigma lobes. Flower colors were based on field observations using the Royal Horticultural Society color chart and label data on herbarium specimens. Capsule lengths were measured from the ovary base to the insertion of calyx/hypanthium at the ovary apex on mature fruits. We denoted capsules as cup- shaped when they are less than twice as long as wide and cylindrical when more than twice as long as wide. We measured four (only for M. holmgreniorum, which is known from few collec- tions) to 36 specimens for each new entity described below, as well as numerous specimens of previously described species. Seed surface characters were assessed using scanning electron microscopy (SEM). Three seeds or more per sampled taxon were examined. Seeds were obtained from herbarium specimens or from field collections, mounted on metal stubs, and coated in gold prior to imaging. Seeds were examined at 20 kV using a Hitachi S-570 SEM, and micrographs were made of two seeds per accession at 700 or 600. Locality data for populations were gathered during fieldwork and from herbarium specimens. Latitude and longitude for new collections were made in the field using GPS. If herbarium specimens lacked field measured coordinates, we used the township, range, and section (TRS) data to infer latitude and longitude coordinates using Graphical Locator (Gustafson 1995). GEOLo- SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA 247 cate v2.13 (www.museum.tulane.edu/geolocate/) and Google Earth (earth.google.com/) were used to estimate latitude/longitude coordinates if only limited locality information was available. Dis- tribution data that could not be georeferenced accurately were not included in distribution maps. To image distributions of species, we imported population locality coordinates into ArcGIS v9.2 (ESRI, Redlands, CA). NEW COMBINATIONS Mentzelia procera (Wooton & Standl.) J. J. Schenk & L. Hufford, stat. et comb. nov. Nuttallia procera Wooton & Standl., Contri- butions from the U.S. National Herbarium, 16: 150, 1913; Mentzelia pumila Torr. & A. Gray var. procera (Wooton & Standl.) J. Darl., Annals of the Missouri Botanical Garden, 21: 169, 1934.—Type: USA, New Mexico, White Sands, 18 August, 1907, Wooton & Standley s.n. (holotype: US; isotype: NMC). Mentzelia lagarosa (K. H. Thorne) J. J. Schenk & L. Hufford, stat. et comb. nov. Mentzelia pumila Torr. & A. Gray var. lagarosa K. H. Thorne, Great Basin Naturalist, 46: 558, 1986. —Type: USA, Utah, Uintah Co., TI1S, R24E, S11, near Watson, Evacuation Creek, 10 mi., 173 degrees from Bonanza, 1708 m, on gravel, 1 August 1980, Goodrich & Atwood 14664 (holotype: BRY; isotype: NY). Mentzelia pumila was treated broadly as a taxon widely distributed in the North American West in the monograph of Loasaceae by Urban and Gilg (1900) and the revision of Mentzelia by Darlington (1934). In contrast, Hill (1977) argued that M. pumila was restricted to the Red Desert of Wyoming, and Holmgren et al. (2005) treated it as restricted to Wyoming and adjacent portions of southern Montana and northeastern Utah, recognizing the name as misapplied to popula- tions outside of this area. Schenk (2009) recog- nized the range of M. pumila as identical to that inferred by Holmgren et al. (2005) and demon- strated that an exemplar for M. pumila from Wyoming is phylogenetically isolated from line- ages referable to M. pumila var. procera (sensu Darlington 1934) from New Mexico and _ sur- rounding areas and M. pumila var. lagarosa (sensu Thorne 1986) from southern Utah and Nevada. The new combinations recognize M. pumila, M. lagarosa, and M. procera as indepen- dent evolutionary lineages. The variety /agarosa was placed as part of M. pumila based on identical chromosome numbers (n = 11) and similar growth habit (Thorne 1986). The pinnatisect laminas and sinuate anticlinal cell walls of seed testal cells of M. /agarosa distinguish it from M. pumila, which has _ pinnately-lobed laminas and straight anticlinal walls on seed testal 248 Fic. 1. cells (Fig. 1). Populations of M. /agarosa occur in Colorado, Nevada, and Utah and are disjunct from the populations of M. pumila. The sister relationship of M. /agarosa remains uncertain (Schenk 2009); however, its narrowly dissected laminas, floral forms, and seed microsculpture characters are similar to taxa such as M. laciniata (Rydb.) J. Darl., M. conspicua T.A. Todsen, and M. filifolia, which is described below. MADRONO [Vol. 57 Scanning electron micrographs of seed coat testal cells. A. Mentzelia pumila, B. M. lagarosa, C. M. procera, D. M. filifolia, E. M. mexicana, F. M. multiflora. Mentzelia procera can be distinguished from M. pumila by sinuate versus straight anticlinal walls of seed coat testal cells (Fig. 1), respectively. The two species have overlapping morphological states, although petal lengths (9.5—-16.3 mm vs. 11.5— 20 mm) and capsule size (9.8—18.8 mm vs. 11.5— 20 mm) are generally smaller in M. procera than in M. pumila. Mentzelia procera occurs in New Mexico and Colorado and is disjunct from M. 2010] Ss Si) 4 1%, } ape FIG. 2. Scale bars = 3 cm. pumila. Molecular phylogenetic results (Schenk 2009) indicate that M. procera is more closely related to M. integra (M. E. Jones) Tidestr. and M. sivinskii (described below) than it is to M. pumila. NEw SPECIES Mentzelia paradoxensis J. J. Schenk & L. Hufford. sp. nov. (Fig. 2A).—Type: USA, Colorado, Montrose Co., Paradox Valley, along Hwy 90, 12.7 rd mi SW of its jct with Hwy 141, E of Bedrock and Dolores River, 38°16.556'N, 108°47.632'W, 2 Jun 2006, L. Hufford 4475 (holotype: WS; isotype: COLO, NY, RM, UC, US). Habitus singularus erectus, axillarus ramus brevus; caudex singularis; folia alterna elliptica vel lanceolata, margine lobata; petala 5, flavidus, spatulata; staminodia extima petaloidea; semi- num testa in alam expansa. Biennial herbs, up to 9 dm tall; taprooted. Main stem erect, straight, lateral branches on distal half of main stem or along the entire main shoot, lateral branches perpendicular to main stem along its basal region but branching acutely upward relative to the main stem along its distal region, branches straight; decumbent branches absent; epidermis pubescent, becoming white, exfoliating with age. Leaves alternate; rosette leaves narrowly to broadly spatulate, petiolate: SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA 249 MAITED STATES 2811700 Type specimens of newly described species. A. M. paradoxensis, B. M. filifolia, and C. M. holmgreniorum. cauline leaves 38-91 X 6-17(21) mm, rachis width 1.6—-4.7 mm; leaves on lower third of main stem oblanceolate, lanceolate, or elliptic, margins dentate to serrate or pinnate with 8—22 lobes, 4.5— 11015) mm apart, lobes nearly opposite, lobe slightly angled towards leaf apex or perpendicular to leaf axis, regular, up to 2—6.8(9) mm long with acute to occasionally rounded apices, margin revolute; leaves on upper third of main stem elliptic to lanceolate with non-clasping bases, margins dentate to serrate or pinnate with 8—16 lobes, 5.4-11.7(114) mm apart, lobes nearly opposite and slightly angled towards leaf apex or perpendicular to leaf axis, regular, up to 2.4— 5(8.5) mm long with acute to occasionally rounded apices, revolute, pubescent, with greater density of simple grappling-hook, complex grap- pling-hook, and needle-like trichomes on abaxial surface, needle-like and occasionally simple grappling-hook trichomes on adaxial side. Inflo- rescence cymose, bract subtending inferior ovary entire, 2.5—11 < 0.4—1.1 mm. Calyx apices acute to attenuate, margin entire, 2.2—-7.6 * 0.8—2.4 mm. Petals five, yellow, pubescent on abaxial surface, narrowly spatulate, 8.3—-15(17.2) x 1.7—-5.3 mm, apex acute to rounded. Androecium yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis papillate or not, anther occasionally twisted or remaining straight following dehiscence; outer whorls of stamens all fertile or fertile and 250 MADRONO New Mexico Fic. 3. [Vol. 57 New Mexico f Arizona Distributions of newly described taxa and selected relatives in USA and Mexico. A. M. longiloba var. yvavapaiensis (#8) and M. longiloba var. longiloba (A). B. Distribution of M. cronquistii (@), M. marginata (fi), and M. paradoxensis (A). C. M. laciniata (A), M. conspicua (@), M. holmgreniorum (+), M. lagarosa (*), and M. filifolia (Wi). D. Distribution of M. sivinskii (@) and M. multiflora (A). E. M. longiloba var. pinacatensis (@). F. M. longiloba var. chihuahuaensis (@ ). staminodial, five outermost stamens in median antesepalous positions petaloid, narrowly spatu- late, 6-13 * 1.2—-3.8 mm, with or without anther, filament or staminode apex rounded to occasion- ally acute; second whorl of stamens all fertile. Ovary inferior, 3-carpellate with 3 placentae, funnelform; style 5.4-10.4 mm long, stigmas three. Fruit a capsule, cup-shaped, 5-9 X 3.7—-6.5 mm, opening apically by three valves, base rounded, no prominent longitudinal costal ridges. Seeds pale grey to light brown with a white to light brown wing, lenticular-ovoid, 1.7—2.7 mm long; testa reticulate, seed coat anticlinal cell walls straight to slightly wavy, central papillae generally 6—11 per cell. Chromosome number not determined. Phenology: Flowering occurs from June to September. Distribution: Populations occur in the Paradox and Gypsum Valleys of western Colorado at 1585-1964 m elevation (Fig. 3). Plants occur on road cuts, valley slopes and bottoms, and sparsely vegetated gypsum knolls. 2010] Etymology: Mentzelia paradoxensis is named for the Paradox Valley of western Colorado. Representative specimens: USA. COLORA- DO. Montrose Co.: W Paradox, Payson 2323 (RM); Paradox, Walker 157 (RM); Paradox Valley, along Hwy 90, 12.5 mi from its jct with Hwy 141, Hufford 4335 (WS); Paradox Valley, along Hwy 90, just E of Bedrock and 0.2 mi E of Dolores River and 1.2 mi W of its jct with River Rd (Rd Y11), Hufford 4336 (WS). San Miguel Co: Hwy 141, on gypsum knoll across from mi post 36, on the western-most gypsum knolls of the E edge of the Big Gypsum Valley, W of Slick Rock, Schenk 972 (WS); Paradox Formation N of mi post 26 on Hwy 141, T46N R16W S14, Atwood 28894 (RM); 19 mi S of 141/145 jct on Hwyl41 between mi post 36-37, Big Gypsum Valley, T43N RI6W S03, Atwood 28897 (RM); ca. 20 mi S of 141/145 jct on Hwy 141, Big Gypsum Valley, 38°01.480'N, 108°38.885'W, Atwood 28899 (RM); T44N RI6W S31, M. Ownbey 1497 (GH, WS); Gypsum Valley, State Hwy 141, 11.6 km (7.2 mi) SW of Basin, T44N RI6W S32, 38°01'32'N, 108°39’01"W, N. & P. Holmgren 13694 (WS). The phylogenetic analysis of Schenk (2009) placed M. paradoxensis as sister to M. marginata (Osterh.) H. J. Thomp. & Prigge. The distribution of M. paradoxensis is south of the range of M. marginata (Fig. 3). Similar to M. cronquistii H. J. Thomp. & Prigge and M. marginata, M. paradox- ensis has trichomes on the abaxial surfaces of petals. Collections of M. paradoxensis have been identified as M. cronquistii, and this may be a consequence not only of the petal trichomes but also their similar leaf laminas that are narrow and have long, acute lobes. Mentzelia paradoxensis differs from M. cronquistii, M. marginata, and other members of section Bartonia in having a shoot system characterized by many short lateral branches, at least in the lower portion of the main stem, that are nearly the same length along the main shoot. The numerous, short branches of M. paradoxensis give the whole shoot a cylindrical form and densely branched appearance. In con- trast, both M. cronquistii and M. marginata have more candelabrum-shaped shoot systems. Ment- zelia paradoxensis further differs from M. cron- quistii in having smaller capsules. In contrast to M. marginata, which has leaf lobe apices that are rounded to occasionally acute, outermost stamens that are fertile, and long capsules (7-14.5 mm), M. paradoxensis 1s characterized by leaf lobes that have acute apices, outermost stamens that are staminodial or fertile, and short capsules (5-9 mm). Mentzelia filifolia J. J. Schenk & L. Hufford, sp. nov. (Fig. 2B).—Type: USA, New Mexico, McKinley Co., W of Gallup, Pima Rd, 2nd rd E of Hilltop Rd, 0.4 miles N of NM Route 264, 1.3 mi E of AZ border, 35°39.126'N, 109°01.571’W, 4 Aug 2006, J. Schenk 1659 SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA 251 (holotype: WS; isotypes: ARIZ, ASC, NMC, NY, RENO, UNM, US, UTC, WS). Habitus singularis erectus; caudex singularis; folia alterna elliptica vel lanceolata, margine pinnatisecta filia; petala 5, flavidus, spatulata:; staminodia extima petaloidea; seminum testa in alam expansa. Biennial herbs, up to 7.5 dm tall; taprooted. Main stem erect, straight, lateral branches on distal half of main stems at acute angles, straight; epidermis pubescent, becoming white, shedding with age. Leaves alternate, rosette leaves un- known; cauline leaves 43—94(115) X< 7.5—27(36) mm, rachis filiform, 1—2.4 mm wide; lower third of main stem oblanceolate to elliptic, margins filiform, pinnatisect with 8—20 lobes, 6-9 mm apart, nearly opposite, perpendicular, regular, up to 3.2—12(15.7) mm long with acute apex, margins revolute; upper leaves oblanceolate to elliptic with non-clasping bases, margins filiform, pinna- tisect with 8—20 lobes, 7-12 mm apart, nearly opposite, perpendicular, regular, up to 5.6—17 mm long with acute apex, margins revolute; pubes- cent, abaxial surface with greater density of simple grappling-hook, complex grappling-hook, and occasionally with needle-like trichomes than adaxial surface, adaxial surface with needle-like trichomes. Inflorescence cymose, bract subtend- ing inferior ovary entire to pinnate, 7—20 = 0.5— 5.6 mm. Calyx 6-11 xX 1-3 mm, apices acute to attenuate, margins entire. Petals five, yellow, glabrous on abaxial surface, oblanceolate, 14— 18.5 xX 3.6-6 mm, apex acute. Androecium yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis not papillate, straight following dehiscence; outer whorl of stamens fertile and staminodial, five outermost stamens in median antesepalous positions petaloid, oblanceolate, 10.3-14(18) & (1.4)2.5-4.4 mm, without anther, staminode apex acute; second whorl of stamens all fertile. Gynoecium 3-carpellate, ovary inferior, funnelform, 3 placentae; style 10—12.5(14) mm long, stigmas 3. Fruit a capsule, cylindrical, 11— 19.3 x 5-7.5 mm, opening apically by three valves, base tapering, costal ridges running lengthwise diminutive. Seeds grey to light brown, lenticular-ovoid, winged, 2.9—3.2 mm; testa retic- ulate, seed coat anticlinal cell walls sinuate, central papillae generally 42-48 per cell. Chro- mosome number 1 = 10 (Thompson 3553 [US]). Phenology: Plants flower from July to August. Distribution: Populations occur in Apache Co., Arizona, and McKinley Co., New Mexico, where they occur on road-cuts and slopes of dark loam and rocky soils at 2122—2133 m elevation (Fig. 3). Etymology: The specific epithet refers to the filiform lobes and narrow rachis of leaf laminas that serve to distinguish M. filifolia from other Mentzelia species of Arizona and New Mexico. ade Representative specimens: USA. ARIZONA. Apache Co.: S of Wheatfields Lake, near turnoff to Crystal, along rd from Lukachukai to Fort Defiance, Mason 2051 (ARIZ); Rte 12, 0.3 mi N of Rte 264, 35°39.792'’N, 109°05.440'W, Schenk 1660, 1661 (WS). NEW MEXICO. McKinley Co.: along rd to Lukachukai, about 3 mi N of Red Lake and 2 mi S of jct of rd to Crystal, H. Thompson 3553 (ARIZ); Gallup, Herrick 893 (US); July 1961 spoils, W. Wagner 161 (UNM); May 1963 spoils, W. Wagner 198 (UNM); near mine entrance along roadside, W. Wagner 314 (UNM); on the S end of the July 1961 spoils, W. Wagner 370 (UNM); N of Gallup, Wooton 2800 (US), 3 Aug 1904, Wooton s.n. (US). Darlington (1934) treated the populations recognized here as M. filifolia as part of M. laciniata. We observe that M. filifolia has leaves that are more filiform than those of M. laciniata and other similar species, including M. conspicua, M. holmgreniorum (described below), and M. lagarosa, which also have thin, pinnate lobes along the narrow rachis of their leaf laminas, but not as narrow as M. filifolia. All five of these species, which occur in the southeastern portion of the Colorado Plateau (Fig. 3), are similar in having yellow petals and seed testal cells that have sinuate anticlinal walls. Henry Thompson recognized this entity earlier by annotating herbarium specimens of M. filifolia using the specific epithet “‘navajoa.”’ Molecular phylogenetic results (Schenk 2009) place M. filifolia as sister to the Chihuahuan Desert endemic M. mexicana H. J. Thomps. & Zavort., but this relationship has little support. Mentzelia filifolia differs from M. mexicana in having pinnatisect versus pinnately-lobed lami- nas, larger flowers, and sinuate versus straight to wavy anticlinal cell walls of the testal cell walls (Fig. 1). We note that the pinnatisect laminas of M. filifolia are morphologically most similar to those of M. Jlaciniata, M. lagarosa, and M. conspicua. The lack of support for the relation- ship of M. filifolia and M. mexicana in conjunc- tion with the morphological similarities it has to other species begs a more thorough study of the phylogenetic relationships of M. filifolia. Mentzelia holmgreniorum J. J. Schenk & L. Hufford, sp. nov. (Fig. 2C).—Type: USA, Arizona, Apache Co., along US Hwy 60 at side rd to Green’s Peak Lookout, 17 mi NW of Springerville, 30°15’N, 109°33’W, 20 Aug 1960, H. Thompson 3108 (holotype: US; isotype: US). Habitus singularis erectus; caudex singularis; folia alterna elliptica vel lanceolata, margine pinna- tisecta; petala 5, flavidus, spatulata; staminodia extima petaloidea; seminum testa in alam expansa. Biennial herbs, up to 5 dm tall; taprooted. Main stem erect, straight, lateral branches on distal half of main stem or along the entire main 2 MADRONO [Vol. 57 shoot, lateral branches at acute upward angles to shoot, curved; epidermis pubescent, becoming white, exfoliating with age. Leaves alternate; rosette leaves narrowly to broadly spatulate, petiolate; cauline leaves 42-89 x 11-31.9 mm, rachis width 2.3—3.6 mm; leaves on lower third of main stem oblanceolate to elliptic, margins pinnatisect with 14—20 lobes, 4.6—-10.3 mm apart, lobes opposite, lobes strongly angled towards leaf apex, regular, up to 4.9-14.4 mm long with rounded apices, margins revolute; leaves on upper third of main stem lanceolate with non- clasping bases, margins pinnatisect with 12-18 lobes, 6.8—7.9 mm apart, lobes opposite, lobes strongly angled towards leaf apex, regular, up to 4.2—-12.4 mm long with rounded or acute apices, margins revolute; pubescent, abaxial surface with greater density of simple grappling-hook, com- plex grappling-hook, and needle-like trichomes than adaxial surface; adaxial surface with simple grappling-hook and needle-like trichomes. Inflo- rescence cymose, bract subtending inferior ovary pinnate, 11.7-19.2 x 2.5-6.4 mm. Calyx 6.5—9.4 x 2—2.7 mm, apices acute to attenuate, margins entire. Petals five, yellow, glabrous on abaxial surface, narrowly spatulate, 13.5—-18.8 x 5.2- 6.6 mm, apex rounded. Androecium yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis not papillate, anther remaining straight following dehiscence; outer whorls of stamens fertile and staminodial, five outermost stamens in median antesepalous positions petaloid, narrowly spatulate, 11.1-16 2.7—5 mm, without anther, staminode apex acute; second whorl of stamens all fertile. Gynoecium 3-carpellate, ovary inferior, funnelform, 3 placentae; style 8.4-10.6 mm long, stigmas three. Fruit a capsule, cylindrical, 13.1— 14.6 * 5.8-6.9 mm, opening apically by three valves, base tapering, no prominent longitudinal costal ridges. Seeds pale gray with a white wing, lenticular-ovoid, 3.7—3.8 mm long; testa reticu- late, seed coat anticlinal cell walls sinuate, central papillae generally 26-51 per cell. Chromosome number n = 10 (Christy 1995). Phenology: Flowering occurs from June to August. Distribution: Populations occur in sandy washes, along roadsides, and disturbed areas in Apache Co., Arizona, at 1493-2225 m elevation (Fig. 3). Etymology: We name M. holmgreniorum to honor Noel and Patricia Holmgren’s contribu- tion to our understanding of Mentzelia and their | work on the flora of the intermountain ‘Vest. Representative specimens: USA. AkKiZONA. Apache Co.: Vernon, Bohrer 1100 (ARL.); Hwy 60, | mi W of Springerville, Dearen 6482 (ARIZ); 16.6 mi SW of Concho, 14 mi E of Showlow along State Hwy 789 & 61, H. Thompson 3215 © (ARIZ); around the headquarters of Canyon de | 2010] Chelly National Monument, TOSN RIOW S22, Halse 250 (ARIZ); 10 mi SE of Springerville, White Mountains, L. Benson 9569 (ARIZ); Greer area, Schmidt 256 (ARIZ); 4 mi E of Mexican Water, Shreve 8981 (ARIZ). Henry Thompson called attention to distinctive collections from Apache Co., Arizona, which he annotated using the nomen nudum “‘showlowen- sis.” Charlotte Christy (1995) also called atten- tion to these populations, which she annotated with the nomen nudem “‘pinkavae.” We agree with Thompson and Christy that these popula- tions are distinct, and this is supported by phylogenetic results (Schenk 2009), in which an exemplar for M. holmgreniorum was recovered in a polytomy that included also the morphologi- cally similar species M. /aciniata, M. conspicua, M. filifolia, and M. lagarosa. Mentzelia holmgre- niorum has shorter petals than M. conspicua, pinnate rather than the entire ovary bracts characteristic of M. laciniata, leaf lobes that are acutely angled toward the leaf apex rather than extending perpendicularly from the axis as characteristic of M. /agarosa, and broader lamina lobes and rachis than M. filifolia. Mentzelia sivinskii J. J. Schenk & L. Hufford. sp. nov. (Fig. 4A). —Type: USA, New Mexico, San Juan Co.: 5 mi N of Bloomfield, 36°46.750'N, 107°58.876'W, 18 July 2005, J. Schenk 1021 (holotype: WS; isotypes: NY, UNM, US). Habitus singularis erectus; caudex singularis; folia alterna angusta elliptica vel lanceolata, mar- gine pinnata; petala 5, flavidus, spatulata; stamino- dia absentia; seminum testa in alam expansa. Biennial herbs, up to 7 dm tall; taprooted. Main stem erect, straight, lateral branches on distal half of main stem or along the entire main shoot, lateral branches at acute upward angles to shoot, straight; epidermis pubescent, becoming white, exfoliating with age. Leaves alternate; rosette leaves narrowly to broadly spatulate, petiolate; cauline leaves 33-112.2 X 2.9-11.4 mm, rachis width 1—2.9 mm; leaves on lower third of main stem oblanceolate to elliptic, margins pinnate with 18—24 lobes, 3.7—9.3 mm apart, lobes opposite and perpendicular to leaf axis, regular, up to 0.8-4 mm long with rounded to acute apices, margins revolute; leaves on upper third of main stem elliptic to lanceolate with non-clasping bases, margins pinnate with 6-16 lobes, 3.1—-12.7 mm apart, lobes opposite and perpendicular to leaf axis, regular, up to 1—5.1 mm long with rounded to acute apices, margins revolute; pubescent, abaxial surface with equal or greater density of simple grappling-hook, complex grappling-hook, and needle-like trichomes than adaxial surface; adaxial surface with needle-like and occasionally simple grappling-hook trichomes. Inflorescence cymose, bract subtending inferior ovary entire, 5—13.4 x 0.4-0.8 mm. Calyx 5.4-9.3 X 1.2-2.9 mm, apices SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA 253 acute to attenuate, margins entire. Petals five, light yellow to yellow, glabrous on abaxial surface, narrowly spatulate, 9-14.7 =< 3.1—6.4 mm, apex rounded. Androecium light yellow to yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis not papillate, anther remaining straight following dehiscence; outer whorls of stamens all fertile, five outermost stamens in median ante- sepalous positions petaloid, narrowly spatulate, 6.3-11.5 2.44.9 mm, with anther occasionally borne on a stalk, filament apex rounded, occa- sionally with notch; second whorl of stamens all fertile. Gynoecium 3-carpellate, ovary inferior, funnelform, 3 placentae; style 4.6-9.9 mm long, stigmas three. Fruit a capsule, cup-shaped, 8.2— 12.7 X 5.1-7.7 mm, opening apically by three valves, base tapering to rounded, no prominent longitudinal costal ridges. Seeds pale gray to light brown with a white wing, lenticular-ovoid, 2.7— 2.8 mm long; testa reticulate, seed coat anticlinal cell walls sinuate, central papillae generally 12-21 per cell. Chromosome number not determined. Phenology: Flowering occurs from June to August. Distribution: Populations are narrowly distrib- uted in San Juan Co., New Mexico, at 1524-1816 m elevation (Fig. 3). Plants occur on knolls, slopes, and roadsides in gypsum or brown clay soils. Etymology: Early collections of M. sivinskii were collected by Robert Sivinski, and we name this entity for his contributions to understanding the flora of New Mexico and the diversity of Mentzelia. Representative specimens: USA. NEW MEX- ICO. San Juan Co.: Jones Mine (abandoned), ca. 2.7 air mi NW of La Plata, 36°58'04’N, 108°12'26"W, Sivinski 6614 (WS); 27 mi S of the CO border on the NM State Hwy 511, Kelley 46 (UNM): on old roadbed across dissected highland bordering canyon, T27N RIOW S18, SW1/4 of NW1/4, Lousre 340 (ARIZ). Schenk (2009) found M. sivinskii to be most closely related to M. integra and M. procera in molecular phylogenetic analyses. Mentzelia sivin- skii is narrowly distributed in San Juan Co., New Mexico, and it overlaps with the northwestern range of M. procera. In contrast, M. integra is distributed in the Great Basin, where it is disjunct from its closest relatives. Although the flowers of these three species are similar, the outermost stamens opposite the sepal lobes are fertile in ©. sivinskii but are staminodial in M. integra and M. procera. The most distinctive features of M. sivinskii compared to its relatives are narrow lobes on leaves and deep sinuses between these lobes, and in these attributes, M. sivinskii converges somewhat on leaf attributes of its geographic neighbor M. /aciniata. Collections of M. sivinskii have been misiden- tified as M. multicaulis (Osterh.) J. Darl. (R. [Vol. 57 ~ MADRONO LZ {\ a ~ < x La o Fic. 4. Type specimens of newly described taxa. A. Mentzelia sivinskii, B. M. longiloba var. chihuahuaensis, C. M. | longiloba var. yavapaiensis, and D. M. longiloba var. pinacatensis. Scale bars = 3 cm. 2010] TABLE |. SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA 255 MORPHOLOGICAL STATES OF THE VARIETIES OF M. LONGILOBA. MAS = median antesepalous stamens, the outermost stamens opposite sepals. All measurements are in millimeters. Character var. longiloba var. chihuahuaensis — var. pinacatensis var. vavapadiensis Leaf length 37-112 35-110 35—110 39-7] Leaf width 8.4-24.6 7.4-27.1 5.5—22.0 8.3-19.] Rachis width 5.32134 2.3—9.0 241753 3.5-6.6 Number of lobes 10—28 12-18 8—50 10-24 Lobe length up to 1.4-6.8 up to 2.3-9 up to 3.3-8.9 up to 2.4-6.5 Petal length 13.3-17.5 11.3-16.3 11.9-19.8 12.6-13.6 Petal width 3.7-6.8 3.1-5.] 4.7-8.9 4.55.6 Petal apex acute to rounded acute to rounded rounded rounded MAS length (6.7)10.6—15.5 11.2-15.4 9.7-16.9 11.4-12.8 MAS width 1.9-5.2 2.44.0 3.3-5.6 2.9-4.2 MAS staminodial yes yes/no yes/no no Capsule length 9.6-16.4 10.0—15.0 7.6-13.2 9.7-15.2 Capsule width 6.0-9.2 5.7-8.3 5.8-8.5 5.7-7.2 Seed length 3.34.0 2.9-3.2 2.9-3.4 3.0—3.4 Anticlinal walls sinuate straight sinuate sinuate Number of papillae 67—106 4-6 26-51 10-21 Distribution California, Arizona, Chihuahua Sonora AZ New Mexico, Texas, Utah, Sonora Sivinski, EMNRD-Forestry Division, personal communication) based on the identification key in Darlington (1934). Mentzelia multicaulis is, however, distributed only in western Colorado and eastern Utah (Holmgren and Holmgren 2002). Unlike the perennial M. multicaulis, which produces multiple aerial branches from a subter- ranean caudex (Holmgren and Holmgren 2002; Schenk and Hufford 2009), M. sivinskii has a single main stem (Fig. 4). NEW VARIETIES OF MENTZELIA LONGILOBA Josephine Darlington (1934) first recognized M. longiloba as a distinct species distributed in eastern Utah and southern California. Although she differentiated M. longiloba from M. multi- flora (Nutt.) A. Gray on the basis of shorter capsules that have acute bases, Felger (1980) treated the two entities as conspecific and recognized M. multiflora subsp. longiloba (J. Darl.) Felger. Our phylogenetic studies indicated M. multiflora s.s. is more closely related to other mentzelias than it is to M. longiloba (Schenk 2009). Mentzelia multiflora s.s. can be distin- guished from M. longiloba by its longer capsules (11.2-26.1 mm versus 7.6—-16.4 mm), attenuate rather than rounded capsule bases, and entire rather than pinnate prophylls. We recognize M. multiflora s.s. as a taxon limited to the eastern side of the Southern Rocky Mountain Front Range, and we present below a new interpreta- tion of the range of M. longiloba. Our phylogenetic analyses identified a set of morphologically and geographically distinct pop- ulations in a polytomy with M. longiloba s:.s. (Schenk 2009). Although these populations can be distinguished from M. Jongiloba s.s. based on micromorphological states of seed coats, they diverge from it otherwise in largely continuous macromorphological states (Table 1). Given the partially continuous morphological variation of these distinctive populations with M. longiloba while having geographical uniqueness, we recog- nize them as varieties of M. longiloba. Our concept of M. longiloba var. longiloha is mostly consistent with that of Darlington’s (1934) M. longiloba, although we recognize additional variation. Based on collections not available to Darlington (1934), we extend the range of ™. longiloba var. longiloba from California and Utah, to include also Arizona, New Mexico, Texas, and northern Mexico. Additionally, we extend the range of morphological variation to recognize longer petals (13.3—-17.5 mm) and longer capsules (9.6—16.4 mm). Among the new varieties of M. longiloba we describe below, var. chihuahuaensis can be distinguished from the others by the straight anticlinal walls and 4-6 papillae on the outer periclinal wall of seed coat testal cells versus the sinuate anticlinal walls and 10-106 papillae per cell among the other varieties (Table 1, Fig. 5). Variety yavapaiensis can be distinguished from var. pinacatensis because all stamens, including the petaloid outermost stamens, are fertile, its petals are shorter, and its leaves have a lower maximum number of lobes (Table 1). We distin- guish var. yavapaiensis from var. /ongiloba primarily by having fewer papillae per outer periclinal wall of seed coat testal cells than the later. The leaves of var. pinacatensis are narrower and its capsules shorter than those of the other varieties (Table 1). 256 FIG, >: MADRONO [Vol. 57 ‘ = 4 Scanning electron micrographs of seed coat testal cells. A. M. longiloba var. chihuahuaensis, B. M. longiloba var. yavapaiensis, C. M. longiloba var. longiloba, and D. M. longiloba var. pinacatensis. Mentzelia longiloba J. Darl. var. chihuahuaensis J.J. Schenk & L. Hufford var. nov. (Fig. 4B). —Type: USA, Texas, Brewster Co., Rte 118, S of 898, on W side of rd with E exposure, 30°05.898'N, 103°35.782'’W, 1385 m elevation, 7 Aug 2004, J. Schenk 901 (holotype: WS; isotypes: ARIZ, NY, UNM, TEX, US). Habitus singularis erectus; caudex singularis; folia alterna elliptica vel lanceolata, margine lobata; petala 5, flavidus, spatulata; staminodia extima petaloidea; seminum testa in alam expansa. Biennial herbs, up to 5 dm tall; taprooted. Main stem erect, straight, lateral branches on distal half of main stem or along the entire main shoot, lateral branches at acute upward angles to shoot, upwardly curved; epidermis, pubescent, becoming white or gray, exfoliating with age. Leaves alternate; rosette leaves narrowly to broadly spatulate, petiolate; cauline leaves 35— 110 X 7.4-27.1 mm, rachis width 2.3-9 mm; leaves on lower third of main stem oblanceolate to elliptic, margins dentate with 12—18 lobes, 8.4— 12.5 mm apart, lobes opposite and perpendicular to leaf axis, regular, up to 2.3-9 mm long with rounded to acute apices, margins revolute; leaves on upper third of main stem lanceolate with non- clasping bases, margins serrate to pinnate with 10 lobes, 6.8-9.4 mm apart, lobes opposite and slightly angled towards leaf apex, regular, up to 2.9-7.5 mm long with rounded to acute apices, margins revolute; pubescent, abaxial surface with greater density of simple grappling-hook, com- plex grappling-hook, and needle-like trichomes than adaxial surface; adaxial surface with simple grappling-hook and needle-like trichomes. Inflo- rescence cymose, bract subtending inferior ovary entire to rarely pinnate, 5.2-9.9 * 0.7—2.6 mm. Calyx 5.5-8.8 xX 1.9-3.3 mm, apices acute to attenuate, margins entire. Petals five, yellow, glabrous on abaxial surface, narrowly spatulate, 11.3-16.3 < 3.1—5.1 mm, apex acute to rounded. Androecium yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis not papillate, anther remaining straight following dehiscence; outer whorls of stamens fertile and stamirodial, five outermost stamens in median antesepalous posi- tions petaloid, narrowly spatulate, 11.2—15.4 x 2.4-4 mm, without anther, staminode apex acute to rounded; second whorl of stamens all fertile. Gynoecium 3-carpellate, ovary inferior, funnel- — 2010] form, 3 placentae; style 6.4-9.8 mm long, stigmas three. Fruit a capsule, cup-shaped, 10-15 x 5.7— 8.3 mm, opening apically by three valves, base tapering, no prominent longitudinal costal ridges. Seeds light brown with a white wing, lenticular- ovoid, 2.9—3.2 mm long; testa reticulate, seed coat anticlinal cell walls straight, central papillae generally 4-6 per cell. Chromosome number not determined. Phenology: Flowering occurs from August to November. Distribution: Populations occur in the Chihua- huan Desert in New Mexico and Texas in the United States and Chihuahua and Coahuila states of northeastern Mexico at 548-1555 m elevation (Fig. 3). Plants occur on sand dunes and along roadsides in dry clay or sandy soils that are often disturbed. Etymology: Mentzelia longiloba var. chihua- huaensis is named for the Chihuahuan Desert, to which it is endemic. Representative specimens: MEXICO. CHI- HUAHUA. Samalayuca Dunes, ca. 5 miles S of Samalayuca, and ca. 35 mi S of Cuidad Juarez, 31°17'N, 106°30'W, Provance et al. 1678 (UCR). COAHUILA. along Rio Grande, just S of Ojo Caliente, river mileage 808.3, 29°11’N, 102°56’W, Hodgson et al. 5265 (UCR). USA. NEW MEXICO. Hidalgo Co.: Taylor Draw at Animas Creek in Upper Animas Valley, 0.4 mi N of the jet with the rd over the mountains to Douglas, on Rte 338, T31S R20W S33, Sanders et al. 3051 (UCR). TEXAS. Brewster Co.: E of Marathon, across from Housetop Mtns on Hwy 90, on S side of rd with N exposure, 30°12.233'N, 102°57.562'W, Schenk 909, 910 (WS): Route 118, at intersection with Calamity Creek Rd, S of Alpine, 30°10.173’N, 103°35.031'W, Schenk 598, 900 (WS); S-facing road-cut on Hwy 90 just E of Alpine (across from stinking cattle feedlot), 30°22.539'N, 103°36.658'W, Hufford 4311 (WS). Jeff Davis Co.: Route 17, S of Boy Scout camp by a few mi, Davis Mountains, N of Fort Davis, 30°48.954'N, 103°45.869'W, Schenk 897 (WS). Terrell Co.: along Hwy 90, E of Sanderson and Dryden, | mi W of Lozier Canyon, 4 mi W of jet with Hwy 1865 (to Pumpville), and ca. 15 mi W of Langtry, 29°54.296'N, 101°49.263'W, Hufford 4312 (WS). Collections of M. longiloba var. chihuahuaensis are often identified as M. multiflora, but their seed coat testal cells can readily distinguish them. Seed coat cells of M. longiloba var. chihuahuaensis have straight anticlinal walls and 4—6 papillae that are centrally located on a raised dome of the outer periclinal wall of each testal cell (Fig. 5), whereas testal cells of M. multiflora have sinuate anticlinal walls and 34-48 papillae per cell _ (Fig. 1). Although the northern portion of the _ range of M. longiloba var. chihuahuaensis extends into southern New Mexico (Fig. 3), it does not SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA Pas overlap with the range of M. multiflora, which reaches it southern limit in northern New Mexico. The range of M. longiloba var. chihua- huaensis marginally overlaps with the southern range of M. procera, which can be differentiated from M. longiloba var. chihuahuaensis by the sinuate anticlinal walls of its seed testal cells (Fig. lc), more narrow leaves (5.3—-14.9 mm vs. 7.4-27.1 mm) with a narrower rachis (1.7—-3.9 mm vs. 2.3-9 mm), a wide leaf base, and greater number of lobes (14-26 vs. 12-18). Mentzelia longiloba J. Darl. var. pinacatensis J. J. Schenk & L. Hufford var. nov. (Fig. 4D). —Type: MEXICO, Sonora, Pinacate Region, 1.1 km N of Pinacate Peak, 31°47’05’N, 113°29'25"W, 950 m elevation, 13 Oct 1986, R. Felger & G. Joseph 86-432 (holotype: ARIZ: isotypes: ARIZ, RSA). Habitus singularis erectus; caudex singularis: folia alterna elliptica vel lanceolata, breva, margine lobata; petala 5, flavidus, spatulata; staminodia extima petaloidea; seminum testa in alam expansa. Biennial herbs, up to 5 dm tall; taprooted. Main stem erect, straight, lateral branches along the entire main shoot, lateral branches at acute upward angles to shoot, upwardly curved: epidermis pubescent, becoming white, exfoliating with age. Leaves alternate; rosette leaves narrow- ly to broadly spatulate, petiolate; cauline leaves 35-110 * 5.5-22 mm, rachis width 2.1—7.3 mm; leaves on lower third of main stem elliptic, margins dentate to pinnate with 18—S0 lobes, 2.9-18.2 mm apart, lobes opposite and perpen- dicular to leaf axis, irregular or regular, up to 3.3-6.9 mm long with acute apices, margins revolute; leaves on upper third of main stem elliptic to lanceolate with non-clasping bases, margins dentate to pinnate with 8—28 lobes, 2.1— 5.5 mm apart, lobes opposite and perpendicular to leaf axis, irregular or regular, up to 1.7—-8.9 mm long with acute apices, margins revolute; pubes- cent, abaxial surface with greater density of simple grappling-hook, complex grappling-hook, and generally with needle-like trichomes than adaxial surface; adaxial surface with simple grappling-hook and needle-like trichomes. Inflo- rescence cymose, bract subtending inferior ovary entire, toothed, or pinnate, 5.4-15.7 x 0.7— 2.8 mm. Calyx 6.7-13.6 X 1.7—-3.5 mm, apices acute to attenuate, margins entire. Petals five, yellow, glabrous on abaxial surface, narrowly spatulate, 11.9-19.8 x 4.7-8.9 mm, apex round- ed. Androecium yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis not papil- late, anther remaining straight following dehis- cence; outer whorls of stamens fertile or stami- nodial, five outermost stamens in median antesepalous positions petaloid, narrowly spatu- 258 MADRONO late, 9.7-16.9 = 3.3—-5.6 mm, with or without anther, filament or staminode apex rounded; second whorl of stamens all fertile. Gynoecium 3- carpellate, ovary inferior, funnelform, 3 placen- tae; style 7.6-11.4 mm, stigmas three. Fruit a capsule, cup-shaped, 7.6—-13.2 x 5.8—-8.5 mm, opening apically by three valves, base rounded, no prominent longitudinal costal ridges. Seeds light brown with a white to light brown wing, lenticular-ovoid, 2.9—3.4 mm long; testa reticu- late, seed coat anticlinal cell walls sinuate, central papillae generally 26-51 per cell. Chromosome number not determined. Phenology: Flowering occurs from March to April. Distribution: Populations are distributed in the Pinacate Desert of Sonora, Mexico, at 200-950 m elevation (Fig. 3). Plants occur on slopes in soils composed largely of decomposed volcanic cinder rocks and ash. Etymology: Mentzelia longiloba var. pinacaten- sis is named for the Pinacate Desert, to which it is restricted. Representative specimens: MEXICO. SO- NORA. Pinacate Region, ash flat adjacent to N end of Mayo lava flow, 2 April 1989, Dimmitt s.n. (ARIZ); ca. 1.8 km NWW of Pinacate Peak, 31°46.5'N, 113°30'W, Felger et al. 19475 (ARIZ); Pinacate Region, ca. 0.5 km W of Campo Rojo (=Red Cone Camp), 31°46’N, 113°27'W, Fe/ger et al. 87-56 (ARIZ); lava flow NE of Crater Elegante, 31°50’30"N, 113°20'W, Fishbein & Meggs 30 (ARIZ); E trail of Pinacate Peak, Sierra Pinacate, NW Sonora, 31°45'N, 113°30'W, 9 Apr 1983, Sherbrooke s.n. (ARIZ); Pinacate Mountains, Red Cone Camp, 31°47'N, 113°27’'W, 19 Mar 1983, Soule s.n. (ARIZ); Sierra del Pinacate, SE of Pinacate Peak, 31.55'N, 113.25'W, Webster 22298 (ARIZ). The Pinacate region of northern Mexico has been shaped by recent volcanic activity during the Pleistocene to Holocene (Ezcurra et al. 1987), and M. longiloba var. pinacatensis is one of several endemic taxa that appear to have evolved on its distinctive soils (Felger 1991). Collections of this taxon have been previously identified as ™. longiloba (or M. multiflora subsp. longiloba). Varity pinacatensis has shorter leaves, more numerous lobes per leaf, narrower petals, petal- oid fertile stamens rather than staminodes in outermost androecial positions opposite sepals, and shorter capsules compared to M. longiloba var. longiloba (Table 1). Mentzelia longiloba J. Darl. var. yavapaiensis J. J. Schenk & L. Hufford var. nov. (Fig. 4C). — Type: USA, Arizona, Yavapai Co., Juniper Mountains, W of Flagstaff, Cross Mountain Rd, near Hwy I-40, 35°11.767'N, 113°18.280'’W, 1576 m elevation, | July 2005, J. Schenk 1011 (holotype: WS; isotype: ARIZ). [Vol. 57 Habitus singularis erectus; caudex singularis; folia alterna elliptica vel lanceolata, margine lobata; petala 5, flavidus, spatulata; staminodia extima petaloidea; seminum testa in alam ex- pansa; chromosoma noven. Biennial herbs, up to 7 dm tall; taprooted. Main stem erect, straight, lateral branches on distal half of main shoot, lateral branches at acute upward angles to shoot, upwardly curved; epidermis pubescent, becoming white, exfoliating with age. Leaves alternate; rosette leaves narrow- ly to broadly spatulate, petiolate; cauline leaves 39-71 X 8.3-19.1 mm, rachis width 3.5—6.6 mm: leaves on lower third of main stem oblanceolate to elliptic, margins pinnate with 14—24 lobes, 3.1— 9.6 mm apart, lobes opposite and slightly angled towards leaf apex, irregular or regular, up to 2.4— 6.5 mm long with rounded or acute apices, margins revolute; leaves on upper third of main stem lanceolate with non-clasping bases, margins pinnate with 10—20 lobes, 5—10.2 mm apart, lobes opposite or alternate and slightly angled towards leaf apex, irregular or regular, up to 2.6—5.8 mm long with rounded or acute apices, margins revolute; pubescent, abaxial surface with greater density of simple grappling-hook, complex grap- pling-hook, and occasionally needle-like tri- chomes than adaxial surface; adaxial surface with simple grappling-hook and needle-like trichomes. Inflorescence cymose, bract subtending inferior ovary entire, 4.6-14.5 0.3—-1.1 mm. Calyx 7.2— 8.1 x 1.9-2.3 mm, apices acute to attenuate, margins entire. Petals five, yellow, glabrous on abaxial surface, narrowly spatulate, 12.6—13.6 x 4.5—-5.6 mm, apex rounded. Androecium yellow, stamens numerous, those of inner whorls shorter than outer whorls, filaments glabrous, anther epidermis not papillate, anther remaining straight following dehiscence; outer whorls of stamens fertile and staminodial, five outermost stamens in | median antesepalous positions petaloid, narrowly spatulate, 11.4-12.8 x 2.9-4.2 mm, without anther, staminode apex acute to rounded; second whorl of stamens all fertile. Gynoecium 3-carpel- late, ovary inferior, funnelform, 3 placentae; style 8.8-10.6 mm long, stigmas three. Fruit a capsule, cup-shaped to cylindrical, 9.7—15.2 * 5.7—7.2 mm, opening apically by three valves, base tapering to rounded, no prominent longitudinal costal ridges. _ Seeds pale gray with a white wing, lenticular- ovoid, 3—3.4 mm long; testa reticulate, seed coat anticlinal cell walls sinuate, central papillae generally 10—21 per cell. Chromosome number 7 | = 9 (H. Thompson 3405 [ARIZ]}). Phenology: Flowering occurs from March to October. | Distribution: Populations are located in) Apache, Coconino, Mohave, Navajo, Pinal, and Yavapai counties in Arizona, where they occur in © sandy washes and along roadsides at 432-1676 m elevation (Fig. 3). 2010] Etymology: Mentzelia longiloba var. yavapaien- sis is named for Yavapai Co., Arizona, where the type specimen was collected. Representative specimens: USA. ARIZONA. Coconino Co.: Wupatki National Monument, Flagstaff, Demaree 43981 (ARIZ). Mohave Co.: 4 mi W of Peach Springs, Kearney & Peebles 12747 (ARIZ); Hualapai Mountains, SE of Hualapai Mountain Park, 35°05’N, 113°52'W, Vasek & Clarke HMS-112 (UCR); Tuweep, rim of Grand Canyon, Cottam 8594 (ARIZ); along main rd between Wolf Hole and Cottonwood Wash, T38N RISW S836, Mason & Phillips 2885 (ARIZ). Navajo Co.: Corduroy Canon, 20 mi SW of Show Low along US Hwy 60, H. Thompson 3218 (ARIZ); State Rte 77, ca. 11 mi S of Navajo Indian Reservation boundary, 3 mi S of Leroux Wash, TION R21E, 23 Sep. 1973, Spaulding s.n. (ARIZ): near Oraibi, 24 July 1958, Haskell & Hevly s.n. (ARIZ); Newberry Mesa N of Winslow, 9 June 1940, Darrow s.n. (ARIZ): Winslow, M. Jones 4112 (ARIZ). Pinal Co.: Casa Grande Ruins National Monument, D. Turner & DeKoker 59 (ARIZ); Pinal Mountains, Kearney et al. 6364 (ARIZ); Sacaton Agency, Gilman 220 (ARIZ); San Pedro Valley, 4.9 mi SE of Main St in San Manuel via rd to San Pedro River, dissected lower Bajada ca. 1 mi W of river, TIOS RI8E, Burgess & Burgess 5950 (ARIZ); W SCHENK AND HUFFORD: TAXONOMIC NOVELTIES IN MENTZELIA 299 of Gila Butte beneath and around new overpass bridge, S. Adams 34 (ARIZ). Yavapai Co.: about 80 mi SE of Kingman, Keaney & Peebles 12586 (ARIZ); Antelope Creek, S Weaver Mountains, Yarnell 7.5 Quad, TION RO4W S19 NW1/4, Butterwick & Hillyard 6871 (ARIZ); Black Hills, 5 mi E of Cherry, 34°35’26’N, 111°59'46’W, Helmkamp 7-17 (UCR); Lynx Lake area of Prescott National Forest, 4.8 mi S of Hwy 69 on 197, ca. 6 mi SE of Prescott, L. & S. Landrum 5591 (UCR); Page Springs, Demaree 44336 (ARIZ); Prescott National Forest, 7.8 mi SW of Prescott on Hwy 89, 34°27’'N, 112°32'W, Vasek & Clarke 66091 1-54 (UCR); SW of Prescott, 2 mi SW of Kirkland Junction, H. Thompson 3405 (ARIZ). Collections of Mentzelia longiloba var. yava- paiensis have often been determined as ™. multiflora, which is consistent with Darlington’s (1934) broad treatment of the latter species. The phylogenetic results of Schenk (2009) demon- strated that Darlington’s (1934) concept of M. multiflora encompassed polyphyletic lineages. Mentzelia longiloba var. yavapdiensis 1s more closely related to M. longiloba var. pinacatensis than it is to M. multiflora s.s. (Schenk 2009). Mentzelia longiloba var. yavapaiensis has short, cup-shaped capsules in contrast to the long, cylindrical capsules of M. multiflora s.s. IDENTIFICATION KEY TO MENTZELIA SECTION BARTONIA FOR TAXA DISCUSSED ABOVE 1. Plants with multiple stems that arise from a subterranean branching caudex, plants often forming rounded UL Seat See ce, Bee ie td DE ot ts Gee ete ae Gee tee oh Wish eaten eee ee pou are eee et Gee nas eon M. multicaulis 1’ Plants with a simple caudex (=single main stem, or multiple stems that arise from a single region) at or above the soil surface 2. Petals pubescent on abaxial surfaces 3. Most leaves above the base of plant shallowly lobed with rounded to acute margins or entire, especially on the secondary and tertiary branches; outer stamens opposite each sepal with anther; w. Bat es MOA te Ht Se Uh anion geet adie fa, anise sei le Sitey M. marginata 3’ Most leaves (except prophylls) pinnately lobed on all orders of branches, lobes acute; outer stamens opposite each sepal with/without anther; w. CO, AZ, NW, UT 4. Lateral branches at acute angles to the main stem and extend to near the distal end of the plant (creating a candelabrum profile with a flat to round top); capsules 6-16 X 5—7.3 mm: AZ, CO, NM, UT i Gia tad, doe Doge Oey, eee a es a ce YG) OS. Ges M. cronquistii 4’ Lateral branches perpendicular or acutely angled to main shoot, generally of nearly equal lengths, lateral branching often dense (creating a cylindrical profile); capsules 5-9 x 3.7-6.5 mm: Montrose and San Miguel counties, CO ... 2’ Petals glabrous on abaxial surfaces 5. Leaves along main stem pinnatisectly lobed On reals 27 imimvOr LONG wi. « ee 6’ Petals less than 26 mm long 7. Petals 13 mm long or less and floral bracts subtending ovaries pinnate.......... Ee ee ee ee ee ee eee M. paradoxensis M. conspicua M. lagarosa 7’ Petals greater than 13 mm long and floral bracts subtending ovary entire to pinnate 8. Lamina lobes filiform, 1.4 mm wide or less, lobes up to 17 mm long.......... M. filifolia 8’ Lamina lobes narrow, greater than | mm wide, lobes up to 14.4 mm long 9. Entire bracts subtending ovaries; leaf lobes slightly angled toward distal tip of leaf or perpendicularto leat axis; NIM and CO: 6 ac. cae fence weet a ia eee os M. laciniata 9’ Pinnate bracts subtending ovaries; leaf lobes strongly angled toward distal tip of leaf; Se aS stead allah bx Bt, Sanne Br ince ee ie 2 M. holmgreniorum 5’ Leaves along main stem entire, dentate, serrate, to pinnately lobed 10. Largest trichomes of leaves with ring-like pedestals of pearly white cells; leaf lobes angled on proximal side of lobe, perpendicular on distal side of lobe; stem epidermis generally glabrous or occasionally pubescent, leaf lobes few, generally less than 12.................... M. integra 260 MADRONO [Vol. 57 10’ Leaf trichomes of leaves without ring-like pedestals of pearly white cells; leaf lobes with isometrically angled sides; stem pubescent, leaf lobes many, generally greater than 6 11. Anticlinal walls of testal cells straight 12. Capsules cup-shaped (less than or equal to twice as long as wide) 13. Capsules 9.6-19 mm long; 4-6 papillae per testal cell ........0.0.00..00.000.. Brule GANT us ae eam ie M. longiloba var. chihuahuaensis 13’ Capsules 5.3-13 mm long; 8—12 papillae per testal cell... .........002.. M. mexicana 12’ Capsules cylindrical (greater than twice as long as wide).................. M. pumila 11’ Anticlinal walls of testal cells sinuate 14. Seed periclinal wall with 67 or more papillae per cell ......... M. longiloba var. longiloba 14’ Seed periclinal wall with 68 or fewer papillae per cell 15. Outermost stamens opposite sepal lobes with anther 16. Leaf rachis 1—2.9 mm wide ee ee RY FR ee Oe Leer elr, wery Ae M. sivinskii M. longiloba var. yavapaiensis 15’ Outer stamens opposite each sepal lobes generally staminodial 17. Capsules 5.3-13 mm long; plants occur on volcanic soils................ eo Re ee eae M. longiloba var. pinacatensis 17’ Capsules 9.6—26 mm long; plants occur on loam soils 18. Upper leaf rachis 1.7—3.9 mm wide; capsules 5.2—7.3 mm wide....... M. procera 18’ Upper leaf rachis 2.1-13.7 mm wide; capsules 5.1-9.2.mm.wide......... ACKNOWLEDGMENTS The authors express their gratitude to R. Sivinski, T. Lowrey, H. Thompson, N. Holmgren, P. Holmgren, and A. Tiehm for sharing their insights on Mentzelia section Bartonia. Funding for this project was provided by the Betty W. Higinbotham Trust, the Rodgers McVaugh Graduate Research Grant from the Ameri- can Society of Plant Taxonomists, the Margaret Williams Research Grant from the Nevada Native Plant Society, the Rexford Daubenmire Grant for Graduate Education, and the Hardman Native Plant Award in Botany. We thank the following herbaria for specimens used in this study: ARIZ, ASC, BM, BYU, CAS, DES, GCNP, GH, ID, JEPS, NMC, NMCR, NY, ORE, OSC, PH, POM, RENO, RM, RSA, TEX, UCLA, UCR, UNLV, UNM, US, UTC, WILLU, and WS. Scanning electron micrographs were imaged at the Franceschi Electron Microscopy Center at Washington State University. We also thank R. Sivinski and N. Holmgren for comments on an earlier version of this manuscript. LITERATURE CITED CHRISTY, C. M. 1995. Systematics of the Mentzelia (section Bartonia) multiflora (Nutt.) A. Gray complex (Loasaceae). Ph.D. dissertation, Arizona State University, Tempe, AZ. DARLINGTON, J. 1934. A monograph of the genus Mentzelia. Annals of the Missouri Botanical Garden 21:103—227. EZCURRA, E., M. EQUIHUA, AND J. LOPEZ-PORTILLO. 1987. The desert vegetation of El Pinacate, Sonora, Mexico. Vegetatio 71:49—60. FELGER, R. S. 1980. Vegetation and flora of the Gran Desierto, Sonora, Mexico. Desert Plants 2:87—114. . 1991. Senecio pinacatensis (Asteraceae): a new species from the Pinacate Region of Sonora Mexico. Phytologia 71:326—332. GUSTAFSON, D. L. 1995. Graphical Locator. Website http://www.esg.montana.edu/gl/ [accessed 19 April 2009]. HILL, R. J. 1977. Variability of soluble seed proteins in populations of Mentzelia L. (Loasaceae) from Wyoming and adjacent states. Bulletin of the Torrey Botanical Club 104:93-101. HOLMGREN, N. H. AND P. K. HOLMGREN. 2002. New mentzelias (Loasaceae) from the Intermountain Region of Western United States. Systematic Botany 27:747—762. ; , AND A. R. CRONQUIST. 2005. Inter- mountain flora, vascular plants of the Intermoun- tain West, USA, vol. 2, part B: subclass Dilleniidae. New York Botanical Garden Press, New York, NY: HUFFORD, L. 2003. Homology and developmental transformation: models for the origins of the staminodes of Loasaceae subfamily Loasoideae. International Journal of Plant Science 164(Suppl.): S409-S439. , M. M. MCMAHON, A. M. SHERWOOD, G. REEVES, AND M. W. CHASE. 2003. The major clades of Loasaceae: phylogenetic analysis using the plastid matK and trnL-trnF regions. American Journal of Botany 90:1215—1228. PRIGGE, B. A. 1986. New species of Mentzelia (Loasaceae) from Grand County, Utah. Great Basin Naturalist 46:361—365. SCHENK, J. J. 2009. A systematic monograph of Mentzelia section Bartonia (Loasaceae): phylogeny, diversity, and divergence times. Ph.D. dissertation, Washington State University, Pullman, WA. , W. HODGSON, AND L. HUFFORD. 2010. A new species of Mentzelia section Bartonia (Loasaceae) from the Grand Canyon, Arizona. Brittonia | 62:1-6. AND L. HUFFORD. 2009. Name changes in the | Mentzelia multicaulis complex (Loasaceae). Novon | 19:117-121. THOMPSON, H. J. AND B. A. PRIGGE. 1986. New species and a new combination of Mentzelia section | Bartonia (Loasaceae) from the Colorado Plateau. Great Basin Naturalist 46:449—-554. | THORNE, K. H. 1986. New variety of Mentzelia pumila | (Loasaceae) from Utah. Great Basin Naturalist | 46:557-558. URBAN, I. AND E. GILG. 1900. Monographia Loasa- — cearum. Nova Acta Leopold 76:1—370. | MADRONO, Vol. 57, No. 4, pp. 261—267, 2010 BRODIAEA MATSONI (ASPARAGACEAE: BRODIAEOIDEAE) A NEW SPECIES FROM SHASTA COUNTY, CALIFORNIA ROBERT E. PRESTON ICF International, 630 K Street, Suite 400, Sacramento, CA 95814 rpreston@icfi.com ABSTRACT A newly recognized endemic species, Brodiaea matsonii, is described. This highly localized species is restricted to a single extended population along Sulphur Creek, in Redding, Shasta County, California. Brodiaea matsonii is a diploid species (1 = 6) closely related to the more widespread B. minor, a polyploid species (7 = 12, 24), from which it differs by the slightly smaller pink flowers and by its habitat parameters. Brodiaea matsonii grows from cracks and crevices in bedrock along an intermittent stream within foothill woodland. Key Words: Asparagaceae, Brodiaea, California, endemism, new species. In 1993, Gary Matson discovered an unusual pink-flowered brodiaea growing along Sulphur Creek, north of Redding, California. Initial attempts to identify the plants suggested a relationship with Brodiaea pallida Hoover, a species known from only three other populations in Calaveras and Tuolumne counties, a disjunc- tion of ca. 335 km. Mr. Matson collected corms from the population and gave them to Dean Taylor, an expert on California’s rare plants. Dr. Taylor, in turn, presented me with a pot of the corms in 2007 at a symposium sponsored by the Northern California Botanists, where I gave a presentation on brodiaeas. In late May, when the plants bloomed, I recognized that they were not B. pallida but morphologically were more similar to B. minor (Benth.) S. Watson. Further investigations of the Sulphur Creek population, including field sur- veys, morphological measurements, and chromo- some counts show that the population is sufficiently distinct to warrant recognition at species rank. TAXONOMIC TREATMENT Brodiaea matsonii R. E. Preston, sp. nov. (Fig. 1).—Type: USA, California, Shasta Co.., Redding, on S side of Keswick Dam Rd, 0.4 mi NE of its junction with Quartz Hill Rd, along W Branch of Sulphur Creek; 40°37'13"N, 122°25'25”"W, elev. 700 ft, 24 Jun 2009, R. E. Preston 2689 (holotype: DAV; isotypes: JEPS, MO, NY, RSA, US). Paratypes: USA, CALIFORNIA. Shasta Co.: Redding, Upper Sulphur Creek, D. W. Taylor s.n. (SJEPS); Redding, along Sulphur Creek, S of Keswick Dam Rd crossing, 04 Jun 2007, R. E. Preston 2548 (DAV); west fork of Sulphur Creek, 04 Jun 2007, R. E. Preston 2547 (JEPS). Differt a B. minor perianthio roseo, costis abaxialibus viridibus, et lobis apicem versus et costis adaxialibus saepe erubescentibus. Corm with coarse fibrous coat, 1-10 cm below ground level; leaves 2—5, subcrescent-shaped in cross-section, less than 15 cm long; peduncle slender, 10—25 cm tall, pedicels less than 36.5 mm long; perianth 17.4—-26.7 mm long, tube urceolate, 6.8—9.4 mm long, lobes ascending, 10.6—17.5 mm long, outer oblong, acute, 3.0—-4.2 mm wide, inner oblanceolate, rounded, 3.6—5.2 mm wide, white to pink, tips and abaxial mid-ribs rose, adaxial mid- ribs green; staminodes 6.2—8.5 mm long, erect and approximate to stamens, lanceolate, white, margin entire, involuted; stamens 4.8—6.4 mm long, filament 2.0—2.8 mm long, tapered to wider base, narrowly winged laterally, anther 4.0— 5.1 mm long, linear, tips of anther lobes erect with V-shaped notch between; ovary obovate, 3.85.6 mm long, style 6.5—9.2 mm long, slightly wider near apex, ovules 5—8 per locule; fruit a loculicidal capsule, ellipsoidal, 5-6 mm long, 3 mm wide, valve apex acute; seeds black, ovoid to rhomboid, finely striate, 1—-1.5 mm long. Chromosome number n = 6 (Fig. 2). The chromosome count was performed on root tip cells from corms collected with R. E. Preston 2548 (DAV) (A. Diebold, University of Missouri- Columbia, personal communication). The species 1s named for its discoverer, Gary Matson (1949-1999), horticulturalist and founder of the Redding Arboretum (Howe et al. 2000). I suggest ““Sulphur Creek brodiaea”’ as the com- mon name. DISTRIBUTION, CONSERVATION, AND ECOLOGY Sulphur Creek brodiaea is restricted to a single occurrence in Shasta Co., California. The species is among the rarest taxa in California, consisting MADRONO FIG. 1. Brodiaea matsonii. a) Lateral view of inflorescence; b) lateral view of flower; c) oblique view of flower; d) top view of flower. FIG. 2. Chromosomes of Brodiaea matsonii from root tip cells (based on R. EL Preston 2548 [DAYV]), 2n (photograph provided by A. Diebold, University of Missouri-Columbia). 2010] TABLE 1. PRESTON: A NEW SPECIES OF BRODIAEA FROM SHASTA COUNTY, CALIFORNIA COMPARISON OF FLORAL CHARACTERS FOR BRODIAEA MINOR, B. NANA, 263 AND B. MATSONII. Measurements were made on fresh material, from one flower per plant, from the type locality of B. matsonii, from 20 populations of B. minor, and from 16 populations of B. nana. Measurements in mm. Brodiaea nana (n = 170) Character mean range Perianth tube WAS: 5.0-9.0 Perianth lobes 14.3 10.0—21.0 Width, inner lobes Se2 4.0—8.0 Width, outer lobes 3.8 3.0—5.0 Staminode 12 6.0—9.0 Filament 1.4 10=2.0 Anther 4.0 3.0—5.0 Stamen 4.4 3.5-5.2 Ovary SE) 2.2—-5.0 Style Bes ANH 7.5 Ovule number DS 12=33 of only a few hundred individuals scattered along a 1.6 km reach of stream channel. The population grows from cracks and crevices in bedrock along the banks and on small rocky islands within the channel of Sulphur Creek, an intermittent stream occurring within foothill woodland. The canopy is characterized by Quercus douglasii Hook. & Arn., Q. wislizeni A. DC., and Pinus sabiniana Douglas, with a shrub understory of Ceanothus cuneatus (Hook.) Nutt., Arctostaphylos viscida C. Parry, and Toxicodendron diversilobum (Torr. & A. Gray) Greene. Associated species include Sidalcea hirsuta A. Gray, Centaurium venustum (A. Gray) Robinson, and Holozonia filipes (Hook. & Arn.) Greene. The Sulphur Creek watershed lies in the region where the Klamath Range, with its predominantly metamorphic geology, intergrades with the predominantly volcanic Cascade Range Foothills. Soils in the area, mapped as Auburn clay loam (Klaseen and Ellison 1974), are formed in material weathered from amphibolite schist, which outcrops extensively along this section of the stream. The elevation ranges from 195 to 215 m above mean sea level. PHENOLOGY Like all brodiaeas (Niehaus 1971), Sulphur Creek brodiaea forms corms that are dormant in the soil during the summer drought. New leaves emerge soon after the start of the rainy season, generally in October or November. Similar to the process described for Triteleia laxa Benth. (Han et al. 1994; Schlising and Chamberlain 2007), the plants spend the next six months or so producing a new main corm and one to many small offsets. Blooming occurs in late May and June, generally two to three weeks later than populations of B. minor in the Redding area. Seed set follows soon after, and all above-ground parts wither and dry during the summer dormant period. Brodiaea minor (n = 204) Brodiaea matsonii (n = 24) mean range mean range 8.6 6.5—11.5 8.0 6.89.4 15.0 9.8-20.5 13.5 10.6—17.5 4.7 3.0—7.0 re 330-56 Be 2.8—5.0 3.6 2.9-4.2 9.1 62 12.5 7.6 6.2585 2.4 14.2 23 2.0—2.8 52 315-7.0 4.5 4.0-5S.1 6.4 4.59.0 5.8 5.0=6.6 4.9 3.2—7.0 4.5 3.5-5.8 8.8 6.0—12.0 7.9 6.5-9.2 173 12—24 20.4 15-24 TAXONOMIC RELATIONSHIPS The genus Brodiaea Smith remains a valuable resource for systematic and ecological investiga- tion, despite having been monographed twice (Hoover 1939; Niehaus 1971). Recent treatments of Brodiaea recognized 14 species and eight subspecies (four species each with two subspecies), most of which are entirely restricted to the California Floristic Province (Niehaus 1971: Keator 1993; Pires 2002). Clarification of some species circumscriptions (Preston 2006a, b) and new species descriptions (Preston 2006b; Chester et al. 2007) have altered these totals, and the forthcoming second edition of The Jepson Man- ual will recognize 18 species, two of which have two subspecies apiece (Pires and Preston in press). The description of Brodiaea matsonii brings the total number of currently recognized Brodiaea taxa to 21, and additional morphometric and phylogenetic studies currently underway are likely to raise that total by several more species. Brodiaea matsonii appears to be most closely related to B. nana Hoover, B. minor, and B. pallida, based on the morphological similarities between them. The flowers of B. matsonii are slightly smaller, on average, than B. minor, and they are somewhat intermediate in size between B. minor and B. nana (Table 1). However, the shape of the stamens and staminodes are closer in all respects to those of B. minor, rather than B. nana or B. pallida (see Figure | in Preston [2006a] for comparison of B. minor and B. nana, and Hoover [1938] for a discussion of B. pallida). The main morphological difference between B. mat- sonii and B. minor is the pink (vs. blue-violet) flowers. Brodiaea species typically have violet flowers, and plants with pink flowers, while not unknown, are unusual. Brodiaea rosea (Greene) Baker was originally recognized and described on the basis of its pink flowers. Brodiaea californica Lindl. has several pink-flowered populations in 264 the Battle Creek/Paynes Creek watershed in Tehama and Shasta counties (Rowntree 1936) and at least two populations of B. sierrae R. E. Preston have individuals with pink flowers (personal observation; G. Hartwell, Paradise, CA, personal communication). Although pink- flowered cultivars of the latter two species have been named (Burbanck 1941), intraspecific taxa have not been proposed for either species on the basis of flower color. All four of these species are characterized by small flowers (generally <2.5 cm long) with a perianth tube narrowed above the ovary. The urceolate perianth tube appears to be a synapo- morphy, as the perianth tube is campanulate or funnel-shaped in all other Brodiaea species. Hoover (1939) proposed a series of intrageneric groups of species he felt were related, based primarily on morphological grounds, and he placed B. nana, B. minor, and B. pallida along with B. stellaris S. Watson in an informal Section **Stellares’’. Niehaus (1971) later added B. insignis (Jeps.) Niehaus to this group. However, B. stellaris and B. insignis lack an urceolate perianth tube and possess other morphological differences that indicate that they are probably not closely related to the other species included in section *“*Stellares”. Reliance on morphological data alone has proved of limited usefulness in resolv- ing relationships between and among Brodiaea species, and further work is needed. Because B. matsonii consists of a_ single population and is morphologically similar to B. minor, what is the basis for recognizing B. matsonii at species rank and not just as a variety or subspecies of B. minor? A review of species circumscription within Brodiaea provides the context needed to justify this decision. Taxonomic circumscriptions within Brodiaea traditionally have been grounded on the mor- phological species concept. Species have generally been distinguished on the basis of discrete differences in the shape of the floral parts, whereas taxa delineated on the basis of size differences or the relative position of floral parts have been treated as varieties or subspecies. Unfortunately, all Brodiaea species are superfi- cially similar, and determining diagnostic char- acters among species based primarily on floral characters can be difficult, especially when using pressed specimens (Smith 1811; Greene 1886; Hoover 1939). Historically, Brodiaea has been much more broadly circumscribed, and the common name “brodiaea”’ is still applied to species now segregated among several genera, including Dichelostemma Kunth and_ Triteleia Dougl. ex Lindl. Although Brodiaea appears closely related to Dichelostemma, Triteleia ap- pears to be only distantly related (Pires and Sytsma 2002). In addition, the presence of umbels, corms, and other morphological similar- MADRONO [Vol. 57 ities between Brodiaea and other lilioid geophytes has made higher order classifications difficult, and Brodiaea has been placed variously in Liliaceae, Amaryllidaceae, Alliaceae, and Themi- daceae (Hoover 1939; Keator 1989; Niehaus 1971, 1980; Fay and Chase 1996). The most recent phylogenetic classification based on mo- lecular data, places Brodiaea and relatives with the Asparagaceae in the subfamily Brodiaeoideae (Chase et al. 2009). Hoover (1939) epitomized the traditional, morphological approach to species circumscrip- tions in Brodiaea. Hoover was familiar with the concepts of ecotypes and ecological plasticity, invoking these ideas to explain some of the intraspecific variation that he observed in Bro- diaea, although he did not apply ecological information to inform his taxonomic treatment. He was also limited by the lack of cytological data for Brodiaea species. Hoover recognized ten Brodiaea species, six of which exhibited minimal intraspecific variation. He reduced four other previously-described species to varieties, citing morphological intermediacy as the basis for his changes in rank. He reduced B. nana to a variety of B. minor, stating that the two taxa intergraded completely. He treated B. leptandra Greene as a variety of B. californica, stating that he could find few morphological differences between the two taxa. He recombined B. terrestris Kellogg as B. coronaria (Salisb.) Engl. var. macropoda (Torr.) Hoover, stating that the primary difference between B. terrestris and B. coronaria was the length of the scapes and pedicels, and that the floral morphology was nearly identical. He treated Brodiaea rosea, a rare serpentine endemic known from only a few locations in Lake, Glenn, and Colusa counties as a variety of B. coronaria, downplaying the morphological differences be- tween them. Hoover also extended the morpho- logical species concept to brodiaeas in the broad sense. He reduced Triteleia modesta H. M. Hall Hoover and TJ. leachiae M. Peck Hoover to varieties (of 7. crocea Greene and T. hendersonii Greene, respectively), noting that the varieties differed morphologically in only minor aspects from the typical forms, except for flower color (Hoover 1941). Hoover later took an approach more in line with the biological species concept (Mayr 1963; Grant 1981), re-elevating B. leptan- dra to species rank on the basis of reproductive isolation, despite its morphological similarity to B. californica (Hoover 1955). He also proposed raising morphologically similar varieties of T. ixioides Greene to species rank on the basis of genetic isolation through ecological and geo- graphic separation (Hoover 1955). Niehaus (1971) employed a multifaceted ap- proach to try to get beyond the morphologically- based species concept that limited Hoover’s understanding of species’ boundaries and the 2010] evolutionary relationships among the species. He attempted to incorporate multiple data sources, including morphology, anatomy, cytology, flavo- noid chemistry, hybridization studies, geography, and ecology, to circumscribe Brodiaea species. Niehaus’ treatment was in accord with the biosystematic species concept (Grant 1981), in which species are defined by reproductive isola- tion as a consequence of genetic or ecological factors, or both. Niehaus agreed with Hoover’s acceptance of B. /eptandra at species rank, citing differences in morphology, chromosome number, range, habitat, and flowering phenology that distinguished it from B. californica. He similarly restored B. coronaria var. macropoda to species rank (B. terrestris) based on morphology, distri- bution, and chromosome number. He also recognized B. nana at species rank, distinguished from B. minor on the basis of chromosome number, distribution, and habitat (albeit as B. minor and B. purdyi Eastw., respectively, as Niehaus had difficulty circumscribing the two species for other reasons [Preston 2006a]). Niehaus was not entirely consistent in his approach and maintained several subspecies despite noting differences in morphology, chro- mosome number, and distribution from the typical subspecies. He maintained B. coronaria subsp. rosea (Greene) Niehaus, although he noted that it was a serpentine endemic with a highly restricted range. He also recognized octoploid (n = 24) populations of B. terrestris as subsp. kernensis (Hoover) Niehaus, despite substantial morphological differences and non-overlapping ranges with hexaploid (7 = 18) subsp. ferrestris. He described polyploid (7 = 20) populations of B. elegans Hoover as subsp. hooveri Niehaus, even though differences in staminode morpholo- gy and distribution distinguish them from n = 8 and n = 16 populations. Following the traditional, morphologically- based species concept, B. matsonii might not warrant recognition at species rank. However, based on the biosystematic species concept, B. matsonii does appear to warrant recognition at species rank. It appears to be reproductively isolated from B. minor by a combination of factors, including chromosome number, habitat preference, flowering phenology, and allopatry. Although B. matsonii is morphologically sim- ilar to B. minor, they have different chromosome numbers. The base chromosome number in Brodiaea is n = 6; the majority of taxa are polyploid with n = 12, 18, and 24, although B. elegans is diploid, having n =8, 16, and 20 cytotypes (Johansen 1932; Burbanck 1941; Nie- haus 1971). Brodiaea matsonii is a diploid, along with seven other species including B. nana and B. pallida. In contrast, B. minor appears to consist of a complex of polyploid populations, with popu- _ lations at lower elevations in the Sierra Nevada PRESTON: A NEW SPECIES OF BRODIAEA FROM SHASTA COUNTY, CALIFORNIA 265 and Cascade Range foothills reported to be octoploid (7 = 24) and populations at higher elevations in the Sierra Nevada reported to be tetraploid (1 = 12) (Niehaus 1971). B. matsonii appears to be genetically isolated from B. minor, therefore, as hybrids between Brodiaea matsonii and B. minor would not be expected to be fertile. Niehaus (1971) found that interspecific hybrids had reduced seed set and reduced pollen fertility, and he stated that few interspecific hybrids were known where Brodiaea species occurred sympat- rically. Traditionally, cytotypes that lack clear differ- ences in morphology, ecology, or distribution have not been recognized as separate taxa, and this approach has also been followed for brodiaeas, both in the narrow and broader sense. Niehaus (1971) acknowledged that several Brodiaea species consisted of cytotypes with two or more ploidy levels but declined to recognize taxa based on chromosome number unless there was a clear morphological boundary between them. Keator (1968) found that Dichelostemma capitatum (Benth.) Alph. Wood consisted of multiple cytotypes but was unable to identify morpholog- ical or ecological differences between them. Similarly, Barkworth (1977) studied polyploidy populations of Triteleia douglasii Watson and found no consistent morphological characteristics that could be used to recognize intraspecific taxa. In the case of B. matsonii, however, flower color and ecology clearly distinguish it from B. minor. Brodiaea matsonii has very different habitat parameters than B. minor. The association of B. matsonii with ephemeral stream habitat is very rare within Brodiaea. Brodiaea pallida similarly occurs along intermittent streams; the population along Sawmill Creek in Calaveras Co. also occurs in blue oak-foothill pine woodland on outcrops of amphibolite schist within the stream channel and along the banks (personal observation). In southern California, B. filifolia S. Watson, which generally grows in grasslands and on vernal pool margins, rarely grows from cracks and crevices in bedrock along stream banks (T. Chester, Fall- brook, CA, personal communication). In con- trast, B. minor occurs across a wide range of habitats, including grassland, vernal pool, seep, meadow, and chaparral, often on substrates of volcanic origin, but also on serpentine and gabbro. However, none of the known popula- tions of B. minor occur along streams; if any do so, they must be extremely uncommon. Popula- tions of B. minor in Redding, some of which are less than a mile from the occurrence of B. matsonil, occur in and adjacent to vernal pools on old alluvial terraces, generally where a hardpan is present. Brodiaea nana is also not known to occur along streams, but like B. matsonii and B. pallida, B. nana is associated with a much narrower range of habitats. It is 266 found primarily in vernal pools, although it also occurs in grassland on thin soil overlying bedrock, where soils become waterlogged follow- ing precipitation. Vernal pools and intermittent streams are both seasonal wetland habitats, having saturated soils during the winter rainy season but drying down during the summer drought. Vernal pools occu- pied by B. nana and B. minor typically dry down by mid-April, whereas the intermittent streams occupied by B. matsonii and B. pallida remain wet until mid-May or later. This extended hydrope- riod is reflected in these species flowering phenol- ogy. Brodiaea nana and B. minor populations that occur 1n vernal pool terrain bloom in April and early May, whereas B. matsonii and B. pallida populations bloom in late May and early June. Although flowering date in Brodiaea species 1s ecologically plastic with respect to annual varia- tion in spring temperatures and rainfall (Niehaus 1971), the later blooming dates of B. nana and B. pallida appears to have a genetic basis as well, because both B. matsonii and B. pallida bloom later than B. nana and B. minor under common garden conditions (personal observation). Recent studies based on molecular data have proved useful for understanding relationships within the Brodiaeoideae and may point a way towards resolving species relationships within Brodiaea (Pires and Sytsma 2002). A phylogenetic analysis of Brodiaea and Dichelostemma based on DNA sequences is currently underway that may help to interpret morphological characters and to identify the origin of polyploid lineages, especial- ly those involving cryptic cytotypes. Preliminary results based on ITS sequences indicate that B. matsonii 1s grouped with a clade that is basal to the clades containing B. minor and B. nana (A. Diebold, University of Missouri-Columbia, per- sonal communication). If that relationship 1s confirmed by the full analysis, then it would add further support to the recognition of B. matsonil at species rank. Key to the Brodiaea Species with the Perianth Narrowed Above the Ovary la. Staminodes erect to spreading, margins strong- ly inrolled; stamens narrowly notched at apex, lacking prominent papillae abaxially; filaments winged laterally, T-shaped in cross-section. 2a. Perianth violet, outer mid-ribs red-violet de feepe INGA SP es Sule ap ice ee Re ret ee Guess B. minor 2b. Perianth pink, outer midribs green, lobe tips and upper inner midribs often rose- PEC cs oot ee oe ee ne B. matsonii Ib. Staminodes erect, margins not to. slightly inrolled; stamens broadly V-shaped at apex, with prominent abaxial papillae; filaments winged abaxially, V- or Y-shaped in cross section. 3a. Perianth lobes paler towards the base; perianth tube slightly narrowed above the MADRONO [Vol. 57 ovary; staminodes as broad as or broader than the outer perianth lobes. ..... B. pallida 3b. Perianth color uniform; perianth tube strongly narrowed above the ovary; stam- inodes narrower than outer perianth NODES aes cate) ee et ere oes ee ets B. nana ACKNOWLEDGMENTS I thank Dean Taylor for providing a pot of plants cultivated from corms collected at Sulphur Creek; April Diebold for performing the chromosome count; Glenn Keator and an anonymous reviewer for constructive comments on the draft manuscript. LITERATURE CITED BARKWORTH, M. E. 1977. Intraspecific variation in Brodiaea douglasii Watson (Liliaceae). Northwest Science 51:79—90. BURBANCK, M. P. 1941. Cytological and taxonomic studies in the genus Brodiaea. Botanical Gazette 103:247-265. CHASE, M. W., J. L. REVEAL, AND M. F. FAy. 2009. A subfamilial classification for the expanded aspar- agalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161:132—136. CHESTER, T., W. ARMSTRONG, AND K. MADORE. 2007. Brodiaea santarosae (Themidaceae), a new rare species from the Santa Rosa Basalt area of the Santa Ana Mountains of southern California. Madrono 54:187—198. FAy, M. F. AND M. W. CHASE. 1996. Resurrection of Themidaceae Salisb. for the Brodiaea alliance, and recircumscription of the Alliaceae, Amaryllidaceae, and Agapanthoideae. Taxon 45:441-451. GRANT, V. 1981. Plant speciation, 2nd ed. Columbia University Press, New York, NY. GREENE, E. L. 1886. Some genera which have been confused under the name Brodiaea. Bulletin of the California Academy of Sciences 2:125—144. HAN, S. S., A. H. HALEvy, R. M. SACHS, AND M. S. REID. 1994. Morphology of flower initiation of brodiaea. Scientia Horticulturae 56:235—243. Hoover, R. F. 1938. New Californian plants. Leaflets of Western Botany 2:128—133. 1939. A revision of the genus Brodiaea. American Midland Naturalist 22:551—574. . 1941. A systematic study of Triteleia. American Midland Naturalist 25:73—100. . 1955. Further observations on Brodiaea and some related genera. Plant Life 11:13—23. Howe, M., D. BURK, AND V. PARKER. 2000. In Memorium — Gary Matson 1949-1999. Fremontia 27:71-72. JOHANSEN, D. A. Californian Liliaceae I. Botany 19:779—783. KEATOR, G. 1968. A taxonomic and ecological study of the genus Dichelostemma (Amaryllidaceae). Ph.D. Dissertation, University of California, Berkeley, CA. . 1989. The brodiaeas. Four Seasons 8:4—11. ———. 1993. Brodiaea. Pp. 1180-1183 in J. C. Hick- man (ed.), The Jepson manual: higher plants of California. 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PRESTON: A NEW SPECIES OF BRODIAEA FROM SHASTA COUNTY, CALIFORNIA 26) AND K. J. SYTSMA. 2002. A_ phylogenetic evaluation of a biosystematic framework: Brodiaea and related petaloid monocots (Themidaceae). American Journal of Botany 89:1342—1359. PRESTON, R. E. 2006a. A reconsideration of Brodiaea minor (Benth.) S. Wats. and Brodiaea purdyi Eastwood (Themidaceae), with the resurrection of Brodiaea nana Hoover. Madrono 53:46—54. . 2006b. Brodiaea sierrae (Themidaceae), a new species from the Sierra Nevada foothills of California, USA. Novon 16:254-259. ROWNTREE, L. 1936. Hardy Californians. The Mac- Millan Company, New York, NY. SCHLISING, R. A. AND S. A. CHAMBERLAIN. 2006. Biology of the geophytic lily, Triteleia laxa (Themidaceae), in grasslands of the northern Sacramento Valley. Madrono 53:321—341. SMITH, J. E. 1811. Characters of a new Liliaceous genus called Brodiaea. Transactions of the Linnean Society 10:1—5. MADRONO, Vol. 57, No. 4, pp. 268—273, 2010 HOWELLANTHUS DALESIANUS, RECOGNITION OF A NEW GENUS AND SPECIES IN TRIBE PHACELIEAE (BORAGINACEAE) GENEVIEVE K. WALDEN' AND ROBERT PATTERSON Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, California 94132 gkwalden@berkeley.edu ABSTRACT Howellanthus (Constance) Walden and R. Patt. is recognized as a new genus, containing a single species H. dalesianus (J. T. Howell) Walden and R. Patt., based on morphological evidence. Howellanthus dalesianus (Boraginaceae) is a small perennial herb, endemic to northern California and restricted to ultramafic soils. The species was first collected in 1936 by Ella Dales Cantelow and Herbert Clair Cantelow, described by John Thomas Howell as Phacelia dalesiana J. T. Howell, and later placed in the monotypic Phacelia subg. Howellanthus by Lincoln Constance. Key Words: Boraginaceae, Howellanthus, Hydrophyllaceae, Hydrophylloideae, Phacelia, Phacelieae, serpentine. Phacelia dalesiana J. TY. Howell was. first described in 1937, and named in honor of Ella Dales Miles Cantelow, a long-time friend and correspondent of John Thomas Howell (Camp- bell n.d.; Howell 1954-1955, 1955, 1955-1957). Ella Dales Cantelow, with her husband, Herbert Clair Cantelow, collected specimens on a trip to the Scott Mountains (Siskiyou Co., California) in 1936. A duplicate was sent to Howell at the California Academy of Sciences for identifica- tion, and the “‘remarkable new phacelia” piqued his interest sufficiently to request an expedition to collect mature fruiting material (Fig. 1) (Cante- low 1937-1940; Howell 1937). The following year Howell and Alice Eastwood traveled to Scott Mountain, guided by the Cantelows, to collect the type specimen. Howell returned again to Scott Mountain in July to collect additional material with mature fruit for the description, and published the new taxon (Howell 1937). Constance (1953) erected the monotypic subg. Howellanthus (n = 8) to contain the distinct species (see Constance 1953, p. 201 for scientific illustra- tion), based upon chromosome number differenc- es and taxonomic revisions of infrageneric divi- sions in Phacelia (Cave and Constance 1942, 1944, 1947, 1950; Constance 1949, 1950, 1951). The subgenus has since remained monotypic in treat- ments of the genus, separated from subg. Phacelia (n = 11) and subg. Cosmanthus (n = 9) (Constance 1963; Constance and Chuang 1982; Halse et al. 1993; Ferguson 1998; Garrison 2007). Phacelia dalesiana has been considered a paleoendemic due to several factors: the system- atic isolation of the taxon within Phacelia; a hypothesized relictual lineage within Hydrophyl- ‘Current address: Department of Integrative Biology, University of California, Berkeley, CA 94720. loideae; and the ecologic specialization on ultramafic substrate, although it lacks the high ploidy level characterizing other paleoendemics (Constance 1953; Favarger and Contandriopou- los 1961; Stebbins and Major 1965). It occurs in the Siskiyou—Trinity mountains, an area of high concentration of Arcto—Tertiary relictual species (Stebbins and Major 1965). The strongest argument to retain the taxon within Phacelia has been common, shared char- acters between species across several infrageneric divisions in the genus: the scorpioid cyme, although few flowered, lax and axillary from a basal rosette in P. dalesiana (Figs. 2 and 3); presence of interstaminal corolla scales (Fig. 4), modified in subg. Cosmanthus or absent in sect. Whitlavia and sect. Gymnobythus; deeply parted style, which can be lobed in some species of the genus; entire leaves and perennial habit; andn = 8, shared with P. stebbinsii Constance and Heckard, P. marcescens Eastw. ex J. F. Macbr., P. glabra Nutt., and P. quickii J. T. Howell. However, these characters are also shared to varying degree with Draperia Torr., Hesperochiron S. Watson, Romanzoffia Cham., and Tricardia Torr. ex S. Watson (Table 1), which left P. dalesiana in an uncertain and isolated relationship with these genera and Phacelia (Walden 2010). The species is easily recognized morphologically as a member of subfamily Hydrophylloideae, but is distinct from other members of tribe Phacelieae and warrants recognition as a separate genus from Phacelia. The taxon can be distinguished from Phacelia by the combination of perennial, acau- lescent habit, axillary few-flowered scorpioid inflorescence, presence of multicellular glandular trichomes with unicellular ellipsoidal heads, deeply parted style, paired pendant ovules, semiorbicular corolla scales, n = 8, and tricolpo- 2010] WALDEN AND PATTERSON: A NEW GENUS IN THE BORAGINACEAE 269 “i BINT. “2 <= = = & ey Fic. 1. Herbert Clair Cantelow, John Thomas Howell and Ella Dales Miles Cantelow on porch at Scott Ranch (1936 photo by grandson Herbert Park Cantelow, courtesy of the Cantelow family and of Ella Dales and Herbert Cantelow papers, California Academy of Sciences Archives). FIG. 2. Habit of Howellanthus dalesianus. Bar = 1 cm. Fic. 3. Close up of corolla, showing semiorbicular scales at base of filaments. Bar = 1 cm. rate pollen with exine striato-rugulate (Constance and Chuang 1982; Di Fulvio and Dottori 1995; Di Fulvio et al. 1999; Walden 2010). TAXONOMIC TREATMENT Howellanthus (Constance) Walden and R. Patt., gen. et stat. nov. Phacelia subg. Howellanthus Constance, Madrono 11:198—203. 1953.—Type species Howellanthus dalesianus (J. T. Howell) Walden and R. Patt., comb. nov., hoc loco designatus. Phacelia dalesiana J. T. Howell, Leaflets of Western Botany, 2:51. 1937.— Type: USA, California, Trinity Co., Summit of Scott Mountain, 25 June 1937, A. Eastwood and J.T. Howell 5014 (holotype: CAS!; isotypes CAS!, UC!, POM!). Genus habitu Hesperochiron, differt cymis laxis paucifloris, vix scorpioideis; trichomata glanduli- fera, stipula multicellularia, capitata ellipsoidea et unicellularia; corolla alba maculis purpureis in fauce; squamae interstaminales semiorbiculares; filamenta glabra basibus non dilatatis; stylus partitus ad basem; capsula suborbicularis, ovar- ium quasi divisum placentis parietalibus spisses- cens, ovulis 2-4, geminatis, pendulis; pollinis grana 3-colporata, striata-rugulata; semina pler- MADRONO [Vol. 57 Fic. 4. Side view of open corolla and unexpanded bud with presence of abundant glandular trichomes. Bar = 1 cm. umque 2, raro 4, alveolata; chromosomatum numerus 7 = 8; species endemica terrae orlundae ex rupibus ultramaficis. Perennial herb, 5—15 cm, densely hairy due to presence of two types of trichomes, unicellular eglandular verrucose-walled conical trichomes, and multicellular glandular trichomes with uni- cellular ellipsoidal heads. Basal leaves rosulate from caudex, cauline leaves alternate, sometimes appearing opposite on stems, reduced upward. Leaf blades oblong to elliptic, simple, margins entire, with lateral incised venation, 10—5O mm long, blades more or less equal to petiole. Inflorescence a lax scorpioid cyme from axils, one- to few-flowered, pedicel 10-20 mm in fruit. Sepals fused at base, lobes 5, unequal, oblanceo- late, accrescent, 3-5 mm long in flower, 4-7 mm long in fruit. Corolla 5-10 mm in diameter, campanulate, fused at base, deciduous, white, sometimes fading lavender in age, throat purple- marked. Corolla scales semiorbicular, fused to base of corolla throat but not to filaments, 2 mm long. Filaments adnate to corolla at base, glabrous, slender, included to slightly exserted, 6-8 mm long, anthers purple, pollen tricolporate, colpi nearly smooth, exine striato-rugulate. Style 2010] WALDEN AND PATTERSON: A NEW GENUS IN THE BORAGINACEAE 271 parted nearly to base, included to shghtly oO 3 3 exserted, 6-7 mm long, glabrous or sometimes is 3 a pubescent—hairy at base. Ovaries one-celled, x g's 8 ovules paired (2-4 per ovary), pendant. Capsule 2/3 "S >e = s 3 4 mm long, subspheric. Seeds 2-4, ovoid, brown, S =f = & 3 S is on =5 = 2.5—-4 mm long, surface alveolate, pubescent. 1 = S 2 S = Oo 6 ge pales f : 8. Blooming May—August. n Gx OQ o0 OH Teo 3 iS DISTRIBUTION a cD) fo 2) = & ee 6 s Howellanthus dalesianus occurs in the ‘Scott S < $ 8 B Fa Mountains and Trinity Alps, near the junction of S Sone 5 . 3 » a Shasta, Trinity, and Siskiyou counties (Fig. 5). 5 2 ao 2a | Ss i322 The type location is Scott Mountain Summit in Sils, 5 S = = es a ae 5 Trinity County, intersected by State Highway 3 3 Cys. Me ae 5 eg Il eee = and the Pacific Crest Trail (Ferlatte 1978). The a Od wm “aus he ts type locality is the most collected single popula- = ~ tion represented in herbaria, due to the accessi- S 3 B pa ae by the vi iroes oe en = oe = orest trails. However, while the type locality ms s 5 5 s , 2 appear accessible today, we speculate that the aes eet. 220 5 a's species was not collected prior to 1936 due to Cl aga = a 6 oes = 05, morphological similarity to Hesperochiron cali- = Bees eo ee Sloe fornicus (Benth.) S. Watson, which also occurs on ee ee Scott Mountain, and the limited presence of Z botanical collectors in the Scott Mountains prior 2 5 to, 1930: aS e o = For three decades Scott Mountain was the only xy S _ # ae 2 known locality for the species, but currently seven ee = +3 oer” z= a populations have been vouchered in herbaria, fz = 6 : eo § 3 Z icc 8 = although three are from single collections. The Z § 63 & SHY 2s USDA National Forest Service Shasta—Trinity c and Klamath have “surveyed and continue to fa on manage fifty populations for the species on public 2 7 3 : ce lands, with additional aaeegnetny reports cs s ° — we @ A. 225 opulations on private land holdings in the rw AS; 3 E 5 = cA 23 6 Z California Department of Fish and Game Nat- a |s Bone os & DE 6 8 ural Diversity Database (Adamson and_Kier- a8 ~ a :3 s S Ss ij os = 2 2 stead-Nelson 1991). The USDA Forest Service a [™\3 Losey eg oka ZeSs lists Phacelia dalesiana as a sensitive species, and = c o = a z a 22 e S , 2 008 be the California Native Plant Society lists Phacelia . OO i SS Rel ok & dalesiana as 4.3 (limited distribution, not very a threatened in California), California Endemic, S = S3.3 (vulnerable) and G3 (vulnerable, no current Sai me 2 3 threats known) (CNPS 2010; Showalter 1991). bh E z S = 3 The elevation of the species ranges from 1600— S s = ed 2 = os = ae m, on pears! flat topography in Oa * I|/6 en =e 2 One or dry meadows in upper montane coniferous 3 2 & E 5 a 8 3 2 e 5 forest (Adamson and Kierstead-Nelson 1991). All z me ome Pisa co ee as z populations are on soils derived from ultrabasic iS S248 92236 ee k < lated with the Trinity Ultramafic sheet = ee a OR oO Sill oS rock associated wi y v cg 082 8S aupuansss (Kruckeberg 1984; Adamson and Kierstead- S : Nelson 1991). Howellanthus dalesianus has been : 2 7 documented on disturbed areas, with ee 4 = litter cover within mature populations, althoug z 3 a = S the species requires bare soil for germination and - s 3 = § e E 6 establishment (Adamson and Kierstead-Nelson 118 22 §& Z ae 1991; Kierstead-Nelson and Engstrom 1991— | VY} 3 5 » § = 2 = E 5 S 5 1993). The species blooms from May to August, ee = és ; 233 BS RS with flowering closely tied to snowmelt for a Ee = &£D 4 VLANnLVAY short season, and fruits until late September. N ~ N Trinity Humboldt : a MADRONO Shasta [Vol. 57 a “ae ae Te a eee see | kc - \ a \ Tehama Fal , oa 100 km Fic. 5. ADDITIONAL SPECIMENS EXAMINED USA. CALIFORNIA. Shasta Co.: Trinity Mountains, saddle between Rattlesnake Hill and Chicken Hawk Hill, at summit on Forest Rd 38N21, at Sardine Spring, 15 June 1993, D. W. Taylor (JEPS). Siskiyou Co.: 4 m NE Scott Mountain, trail from Robbers Meadows to Kangaroo Lake, 1 June 1977, S. Horner 110 (JEPS); Mt. Shasta Ranger District, Mumbo Basin at 40N26 and 38N24, 20 June 1991, J. Kierstead 91—68 (ST); Scott Mountain divide, 17 May 1947, H. L. Mason 14773 (UC); Kangaroo Lake Campground, along streams, 25 July 1969, F. W. Oettinger s.n. (CAS, RSA, UC); Kangaroo Lake, 25 July 1967, D. Parker and W. Roderick s.n. (CAS); Scott Mountain Summit, 21 June 2005, R. W. Patterson & S. Santos 1982 (SFSU); Scott Mountain rd summit, upper end of N meadow, 14 June 1963, W. Roderick s.n. (JEPS); Scott Mountain campground, 18 June 2010, G. K. Walden 324 (SFSU); Scott Mountain camp- ground, 7 July 2010, G. K. Walden 332 (SFSU). Trinity Co.: Summit of Scott Mountain, 21 May 1936, E. D. Cantelow 1276 (CAS); Summit of Scott Mountain, 9 June 1939, E. D. Cantelow 2891 (CAS); Summit of Scott Mountain, 23 June 1948, E. D. Cantelow s.n. (CAS); Near Scott Mountain Summit, close to the town of Callahan, Distribution of vouchered collections of Howellanthus dalesianus in Siskiyou, Trinity, and Shasta counties, California. Starred locality indicates type locality on map. China Mountain quadrangle 1 July 1978, Clifton and Ground 1662 (UC); Summit of rd over Scott Mountain, sandy flat of rocky meadow, | June 1946, L.Constance and R. H. Shan 3070 (CAS, UC); Trail from Deer Flat to Shiny Lake ca. 1/4 m above Deer Creek Flat, 8 July 1976, J. DiTomaso 600 (UC); Along USFS trail 8W13 from Deer Flat to Shimmy Lake, 1/4 to 1/2 m E of Deer Flat Camp, 7 July 1976, W. J. Ferlatte and J. DiTomaso 1776 (CAS, JEPS); Summit of Scott Mountain, N of Carrville, 24 August 1936, J. T. Howell 12736 (CAS); Summit of Scott Mountain, N of Carrville, 30 July 1937, J. T. Howell 13691 (CAS); Scott Mountain summit on rd from Carrville to Callahan, 20 May 1980, J. T. Howell, T. C. Fuller and G. D. Barbe 53545 (CAS); Summit of Scott Mountain, rocky ser- pentine soil, 9 June 1958, D. Parker 649 (CAS, RSA); Scorpion Lake, Trinity Mountains, west- erly upper flank Bonanza King, 13 June 2002, D. W. Taylor 18156 (JEPS). Shasta Co.: Trinity Mountains, saddle between Rattlesnake Hill and Chicken Hawk Hill, at summit on Forest Rd 38N21, at Sardine Spring, 15 June 1993, D. W. Taylor 13656 (JEPS). ACKNOWLEDGMENTS This represents a portion of work resulting from a master’s thesis by GK W, and was previously presented, 2010] in part, by the authors at the California Botanical Society Graduate Student Meeting 2010, San Jose State University, California, as “*Phacelia dalesiana J.T.How- ell, a remarkable paleoendemic phacelia.”’ This project was supported, in part, by funding to GKW from NSF GRFP, NSF TREE, and NSF GK12 fellowships and research grants by SFSU Department of Biology and CNPS chapters. We are grateful to the following for field, herbarium, archive, and laboratory assistance: Deb Trock, Andrew Doran, Ellen Dean, Sula Vanderplank, Danielle Castronovo, the Cantelow family, Frank Cipriano, Dennis Desjardin, Julie Kier- stead—Nelson, Jim Linnberg, and Trigger (service dog of GKW). We thank the editor and two anonymous reviewers who substantially improved this paper. LITERATURE CITED ADAMSON, B. AND J. KIERSTEAD-NELSON. 1991. Guidelines for identifying suitable habitat for Phacelia dalesiana within its range. Unpublished report for USDA, Forest Service, Redding, CA. CAMPBELL, M. W. n.d. Cantelow biography, Ella Dales and Herbert Cantelow papers. Box 1, folder 1. Archives, California Academy of Sciences, San Francisco, CA. CANTELOW, E. D. M. 1937-1940. Field notebook. Ella Dales and Herbert Cantelow papers. Box 3, folder 1. Archives, California Academy of Sciences, San Francisco, CA. CAVE, M. S. AND L. CONSTANCE. 1942. Chromosome numbers in the Hydrophyllaceae. University of California Publications in Botany 18:205—216. AND . 1944. Chromosome numbers in the Hydrophyllaceae, I]. University of California Publications in Botany 18:293—298. AND . 1947. Chromosome numbers in the Hydrophyllaceae, III. University of California Publications in Botany 18:449—46S. AND . 1950. Chromosome numbers in the Hydrophyllaceae, IV. University of California Publications in Botany 23:363—382. CALIFORNIA NATIVE PLANT SOCIETY (CNPS). 2010, Inventory of rare, threatened, and endangered plants of California, 8th ed. California Native Plant Society, Sacamento, CA, Website http://www. rareplants.cnps.org/ [accessed 5 January 2010]. CONSTANCE, L. 1949. A revision of Phacelia subgenus Cosmanthus (Hydrophyllaceae). Contributions to the Gray Herbarium 168:1-48. 1950. Some interspecific relationships in Phacelia subgenus Cosmanthus. Proceedings of the American Academy of the Arts 78:135—147. . 1951. Hydrophyllaceae. Pp. 476-532 in L. R. Abrams (ed.), Hlustrated flora of the Pacific states, vol. 3. Stanford University Press, Stanford, CA. . 1953. Howellanthus, a new subgenus of Phacelia. Madrono 11:198—203. . 1963. Chromosome number and classification in Hydrophyllaceae. Brittonia 15:273—285. AND T.-I. CHUANG. 1982. SEM survey of pollen morphology and classification in Hydrophyllaceae (Waterleaf Family). American Journal of Botany 69:40-S3. WALDEN AND PATTERSON: A NEW GENUS IN THE BORAGINACEAE 213 D1 FuULvio, T. E. AND N. DOTTORI. 1995. Contribucion al conocimiento de tricomas y emergencias en Hydrophyllaceae. Clasificacion y consideraceones taxonomicas. Kurtziana 24:19—24. M. T. CosA, AND N. DoTToORI. 1999. Morfologia y vascularizacion floral de Draperia, Emmenanthe, Hesperochiron, Romanzoffia, y Tri- cardia (Phacelieae, Hydrophyllaceae). Kurtziana 27:187-209. FAVARGER, C. AND J. CONTANDRIOPOULOS. 1961. Essai sur l’endémisme. Bulletin of the Society Botany Suisse 71:383—408. FERGUSON, D. M. 1998. Phylogenetic analysis and relationships in Hydrophyllaceae based on ndhF sequence data. Systematic Botany 23:253—268. FERLATTE, W. J. 1978. Notes on two rare, endemic species from the Klamath region of northern California, Phacelia dalesiana (Hydrophyllaceae) and Raillardella pringlei (Compositae). Madrono 25.158. GARRISON, L. M. 2007. Phylogenetic relationships in Phacelia (Boraginaceae) inferred from nr/ITS' se- quence data. M.S. thesis. San Francisco State University, San Francisco, CA. HALSE, R. R., D. WILKEN, AND R. PATTERSON. 1993. Hydrophyllaceae. Pp. 691-706 in J. C. Hickman (ed.), The Jepson manual: higher plants of Califor- nia. University of California Press, Berkeley, CA. HoOwEeELL, J. T. 1937. A remarkable new Phacelia. Leaflets of Western Botany 2:51—52. . 1954-1955. Correspondence to Mrs. Cantelow. Ella Dales and Herbert Cantelow papers. Box 3, folder 5, book 1. Archives, California Academy of Sciences, San Francisco, CA. . 1955. Correspondence to Mrs. Cantelow. Ella Dales and Herbert Cantelow papers. Box 3, folder 5, book 2. Archives, California Academy of Sciences, San Francisco, CA. . 1955-1957. Correspondence to Mrs. Cantelow. Ella Dales and Herbert Cantelow papers. Box 3, folder 5, book 3. Archives, California Academy of Sciences, San Francisco, CA. KIERSTEAD-NELSON, J. AND T. ENGSTROM. 1991— 1993. Phacelia dalesiana — three year disturbance response study. Unpublished report for the USDA, Forest Service, Redding, CA. KRUCKEBERG, A. R. 1984. California serpentines: flora, vegetation, geology, soils, and management problems. University of California Publications in Botany 78:1—180. SHOWALTER, K. (ed.). 1991. Decision memorandum. Study of response to disturbance by Scott Moun- tain Phacelia, a sensitive species. Mt. Shasta Ranger District, Siskiyou County. USDA Forest Service, Redding, CA. STEBBINS, G. L. AND J. MAJOR. 1965. Endemism and speciation in the California flora. Ecological Monographs 35:1-—35. WALDEN, G. K. 2010. Phylogeny of infrageneric relationships within Phacelia (Boraginaceae) in- ferred from chloroplast sequence data. M.S. thesis. San Francisco State University, San Francisco, CA. MADRONO, Vol. 57, No. 4, pp. 274-275, 2010 NOTEWORTHY COLLECTIONS CALIFORNIA ACACIA DEALBATA Link. (FABACEAE).—Shasta Co., City of Redding, on a large mid-channel gravel bar in the Sacramento River approximately 0.40 km S of the Bonneyview Bridge, associated species include Populus fremontii, Quercus lobata, Robinia pseudoacacia, Salix lucida subsp. lasiandra, Salix exigua, Rubus discolor, Nerium oleander, Cytisus scoparius, Equisetum arvense, Carex barbarae, and Cynodon dactylon, Enter- prise USGS 7.5’ quadrangle, T31N R4W SE” Sec. 18, UTM 10T 0554595E 4487377N, elev. 134 m, 1 December 2010, L. Lindstrand IIT, s.n. (North State Resources Herbarium! [private], CDA); Shasta Co., on upland bluff 0.48 km below Shasta Dam on E side of Sacramento River, associated species include Ailanthus altissima, Quercus kelloggii, Pinus sabiniana, Robinia pseudoacacia, Arctostaphylos viscida, Cercis occidentalis, Eriodictyon californicum, Heteromeles arbutifolia, and Cytisus scoparius, also observed in nearby riparian habitat adjacent to Sacramento River associated with Salix lucida subsp. lasiandra, Salix exigua, Nerium oleander, Cephalanthus occidentalis, Fraxinus latifolia, Alnus rhombifolia, Vitis californica, and Rubus discolor, Shasta Dam USGS 7.5’ quadrangle, T33N RSW NE'”% of SW'% Sec. 15, UTM 10T 0548826E 4507166N, elev. 232 m, 7 December 2010, L. Lindstrand I, s.n. (North State Resources Herbarium!’ [private], CDA); Shasta Co., City of Redding, on a mid-channel gravel bar in the Sacramento River approximately 0.32 km S of the Highway 44 Bridge, associated species include Quercus lobata, Robinia pseudoacacia, Salix lucida subsp. lasian- dra, Salix exigua, Rubus discolor, Equisetum arvense, Carex barbarae, and Juncus sp., Enterprise USGS 7.5’ quadrangle, T32N R4W NE” of SW'% Sec. 31, UTM LOT 0553553E 4492622N, elevation 140 m, 7 December 2010, L. Lindstrand III, s.n. (North State Resources Herbarium! [private], CDA); Shasta Co., City of Redding, along rocky western shoreline of the Sacra- mento River approximately 0.80 km S of Keswick Dam, associated species include Ai/anthus altissima, Quercus wislizenii, Robinia pseudoacacia, Salix lucida subsp. lasiandra, Salix exigua, Nerium oleander, Cytisus sco- parius, Arctostaphylos viscida, Cercis occidentalis, Erio- dictyon californicum, Ceanothus cuneatus, and Brickellia sp., Redding USGS 7.5’ quadrangle, T31N RSW NW'%4 of NW'%4 Sec. 28, UTM 10T 0546918E 449S5017N, elevation 152 m, 7 December 2010, L. Lindstrand ITT, s.n. (North State Resources Herbarium! [private], CDA). Previous knowledge. Acacia dealbata is native to southeastern Australia and an invasive species to California. In California the species is known to occur in the western North Coast Ranges, San Francisco Bay Region, western South Coast Ranges, and the South Coast (Hickman 1993; DiTomaso and Healy 2007). The species has also been recorded in the northern Sierra Nevada Foothills in Butte Co. (Calflora 2010). These findings represent the first records of Acacia dealbata in Shasta Co., and the northernmost-recorded extent of 'North State Resources, Inc. Herbarium, 5000 Bechelli Lane, Suite 203, Redding, CA 96002. the species in interior northern California. At the Shasta Dam site the species was first observed on 4 November 2010, when plant material was collected in the field and given a tentative identification of an unknown Acacia. Following further detailed examination, the plant was identified as Acacia dealbata. Additional plant material was collected from the site on 7 December 2010. Plant material was collected from the City of Redding sites on | and 7 December 2010 when the locations were first observed. Vouchers from all locations were sent to the California Department of Food and Agricul- ture Plant Pests Diagnostics Center for annotation, where Botany Laboratory staff confirmed the species identification. Significance. Acacia dealbata is included in the 2007 California Invasive Plant Council Invasive Plant Inventory and assigned a ‘‘Moderate”’ rating (Cal-IPC 2006). These findings represent the first recorded observations of Acacia dealbata in Shasta Co., and significant (between approximately 160 to 180 km) northern extensions of the known species range in interior California. Multiple age classes, fruiting indi- viduals, and seedlings, were observed at all locations, suggesting these populations reproduce and are capable of expansion. —LEN LINDSTRAND III, Fisheries/Wildlife Biologist, Terrestrial Biology Program Manager, North State Resources, Inc., 5000 Bechelli Lane, Suite 203, Red- ding, CA 96002. lindstrand@nsrnet.com. OREGON MATTHIOLA INCANA (L.) W. T. Aiton (BRASSICA- CEAE).—Clatsop Co., N end of Arch Cape on sand- stone cliff overlooking the Pacific Ocean, 45.82333°N, 123.96233°W, elev. 5 m, with Holcus lanatus, Gaultheria shallon, Rubus, Fragaria, Lonicera, Festuca, Aira, 16 September 2008, R. R. Halse 7553 (OSC, MO). Previous knowledge. commonly cultivated ornamental. It has become naturalized in California and Texas on ocean cliffs and bluffs and sandy areas (Al-Shehbaz 2010.) Significance. First report for Oregon. BRASSICA OLERACEA L. (BRASSICACEAE).—Lane Co., along U.S. Hwy. 101 ca. 8.4 m S of Yachats, 44.20830°N, 124.11411°W, elev. 15 m, coastal bluff with | Marah, Rubus, Equisetum, Vicia, Gaultheria, Heracleum, 25 May 2010, R. R. Halse 7880 (OSC, duplicates to be | distributed). Previous knowledge. This native of Europe is a | commonly cultivated vegetable crop. It has become naturalized in coastal California and in the northeastern U.S. and adjacent Canada (Warwick 2010). Significance. First report for Oregon. SCHOENOPLECTUS CALIFORNICUS (C. A. Mey.) Sojak (CYPERACEAE).—Lane Co., Siuslaw National | Forest, Baker Beach area ca. 7 m N of Florence, 44.09333°N, 124.11920°W, elev. 5 m, very common in This European native is a. 2010] shallow waters around the edge of Lily Lake with Ulex, Rubus, Lysichiton, Callitriche, Oenanthe, Spiraea, stems triangular, to 3 meters tall, 9 July 2009 R. R. Halse 7719 (OSC, RSA, WTU, NY). Previous knowledge. This native species 1s found in marshes from California eastward across the southern U.S. to North Carolina (Smith 2002). Significance. First report for Oregon. ORNITHOPUS PERPUSILLUS L. (FABACEAE).—Lane Co., Washburne State Park, N of Hecata Head, bluff above the beach, on the trail by the picnic area, in Pinus contorta woods, common, T16S, R12W, Sec. 22, 7 June 2004, K. L. Chambers 6398, determined by R. R. Halse in 2009 (OSC); same area, Carl G. Washburne Memorial State Park off of U.S. Hwy. 101 ca. 12 m N of Florence, 44.16110°N, 124.11708°W, elev. 14 m, weed around edges of trails with Picea, Vaccinium, Maianthemum, Gaultheria, 9 July 2009, R. R. Halse 7723 (OSC, WTU, NY, RSA, VDB); same location, common weed around parking areas with Trifolium spp., Lolium, Medicago, Vicia, 25 May 2010, R. R. Halse 7876 (OSC, MU, duplicates to be distributed). Previous knowledge. This native of Europe is known from Pennsylvania (Rhoades and Klein 1993). Significance. First report for Oregon. TRIFOLIUM RETUSUM L. (FABACEAE).—Jackson Co., along State Hwy. 62 near its junction with Corey Road, southern edge of White City, 42.41491°N, 122.85540°W, elev. 398 m, weedy roadside with Matricaria, Erodium, Trifolium spp., Hordeum, Vicia, Poa, 31 May 2010, R. R. Halse 7906 (OSC, MU, duplicates to be distributed). Previous knowledge. This European native is known from California (Hrusa et al. 2002). Significance. First report for Oregon. —RICHARD R. HALSE, Department of Botany and Plant Pathology, 2082 Cordley Hall, Oregon NOTEWORTHY COLLECTIONS 275 State University, Corvallis, OR, 97331. halser@science. oregonstate.edu. LITERATURE CITED AL-SHEHBAZ, I. A. 2010. Matthiola. Pp. 253-255 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, vol. 7. Oxford University Press, New York, NY. CAL-IPC. 2006-2007, California invasive plant inven- tory, and update. Cal-IPC Publication 2006-02. California Invasive Plant Council, Berkeley, CA. Website: http://www.cal-ipc.org/ip/inventory/index. php [accessed 10 November 2010]. CALFLORA. 2010, The Calflora database. Berkeley, CA. Website: http://www.calflora.org/ [accessed 10 November 2010]. DITOMASO, J. M. AND E. A. HEALY. 2007. Weeds of California and other western states. Division of Agriculture and Natural Resources, University of California, Berkeley, CA. HICKMAN, J. C. (ed.) 1993, The Jepson manual: higher plants of California. University of California, Berkeley, CA. HRUSA, F.; B. ERTTER, A. SANDERS, G. LEPPIG, AND E. DEAN. 2002. Catalogue of non-native vascular plants occurring spontaneously in California be- yond those addressed in The Jepson Manual part I. Madrono 49:61—98. RHOADES, A. F. AND W. M. KLEIN, JR. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. American Philosophical Society, Phila- delphia, PA. SMITH, S. G. 2002. Schoenoplectus. Pp. 44-60 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, vol. 23. Oxford University Press, New York, NY. WARWICK, S. I. 2010. Brassica. Pp. 419-424 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, vol. 7. Oxford University Press, New York, NY. MADRONO, Vol. 57, No. 4, p. 276, 2010 PRESIDENT’S REPORT FOR VOLUME 57 Dear CBS member, The past year has been an exciting time for the Council of the California Botanical Society. The Council this year has moved closer to bringing the Society totally online through the website. The website has been redesigned to update approaches and bring more information to the membership. Please check it out and give us ideas for making it better for you. We have made great progress with Madrono thanks to the editors; submission of manuscripts and reviews have now been converted to an online process. Meanwhile, we have a proposal to get all the back issues of Madrono available through JSTOR. This year, thanks to the Treasurer, we also established a Finance Subcommittee, staffed by our Treasurer, the Past- President and President. This permits us to establish financial investment goals that might grow our current funds and help us keep future cost increases minimal to the membership. Given that the California Botanical Society was established in 1913, this upcoming year will represent the 98th year of the Society. Yes, that means we are already planning for the Centennial Celebration of our Society. We have negotiated with the California Native Plant Society to join them in their large meeting in January 2012 and provide the Banquet Speaker for the event. We are looking at 2012 as the initiation of a year of field trips sponsored to some extent by the Society, perhaps some trips reenacting historic “‘phyto-jogs” in California’s past. Our membership base is the foundation of the Society and your support allows us to promote botanical research and education. This year, in addition to encouraging other botanists you know to join the Society, we are also hoping to hear from you about your ideas for the celebration of our 100th year. Let us know by emailing or writing to any member of the Council. We’re certain that you harbor some incredible ideas for the Centennial! Increasing our membership is always a priority, so please continue to encourage your colleagues to join us and to publish in Madrono. This is especially true of our younger colleagues; as we move online we hope to be more attractive to the younger cohorts of botanists raised in an all-online computer age. Also, please consider providing a sponsoring membership or subscription to a foreign scientist or scientific institution to support botanical research in economically depressed, developing countries. For more information on making such a gift, please con- tact Corresponding Secretary Heather Driscoll (hdriscoll@ berkeley.edu). The Society also welcomes gifts or other contributions to our endowment. V. Thomas Parker December 2010 MADRONO, Vol. 57, No. 4, p. 277, 2010 EDITORS’ REPORT FOR VOLUME 57 We are pleased to report the publication of this volume of Madrono by the California Botanical Society (CBS) in 2010. The journal is continuing to reduce turnaround time, as we now average six months between submission and publication. With the new electronic submission and manuscript review process, we feel we are well on our way to a much smoother review process. As always, we are extremely grateful to all the individuals who serve are reviewers and contribute to the quality of the journal. This year we received 26 new manuscripts and 24 were accepted for publication. Several manuscripts were also carried over from the previous year. The current volume includes articles (including Notes), new taxa, Noteworthy Collections, and Book Reviews. There was a mix of systematic and ecological manuscripts submitted. Many of the systematic manuscripts incorporated current molecular techniques and data as well as cutting edge data analysis methods. It is notable that manuscripts reporting taxa new to science are being submitted on a regular basis. As Editors, we have enjoyed our interactions with contributors and reviewers this past year and look forward to another year of continuing the long and distinguished tradition in botanical publication represented by Madrono. Tim Lowrey Richard Whitkus December 2010 MADRONO, Vol. 57, No. 4, p. 278, 2010 David Ackerly Patrick Alexander Wendy Applequist Debra Ayres Tina Ayers Donovan Bailey Brent Blair Gregory Brown Anita Cholewa Laurie Consaul Stephen Davis Steven Downie Taly Drezner Donna Ford-Wertz Daniel Harder Ronald Hartman Harmut Hilger Noel Holmgren Philip Jenkins Richard Jensen Sophie Karrenberg Glenn Keator Peter Kevan Leah Larkin Joshua Leffler J. David Ligon Barnery Lipscomb Diane Marshall Susan Mazer Joseph McBride REVIEWERS OF MADRONO MANUSCRIPTS 2010 Niall McCarten Douglas McCreary Charles McDonald Leslie McFadden Francisco Molina Freaner Esteban Muldavin David Oline Elizabeth Painter Ingrid Parker William Parker Robert Patterson Gitte Petersen J. Chris Pires William Pockman Robert Preston Barry Prigge Nishanta Rajakaruna Joanna Redfern Lauren Ruane Bjorn Salomon Michael Simpson Robert Sivinski John Strother Lynn Sweet Detlev Vogler Mark Welch Dieter Wilken Carol Wilson Blair Wolf MADRONO, Vol. 57, No. 4, pp. 279-280, 2010 INDEX TO VOLUME 57 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. Abies magnifica var. critchfieldii, new var. from Sierra Nevada, CA, 141. Acacia dealbata, noteworthy collection from CA, 274. AFLP (see Pyrrocoma) Allium peninsulare, noteworthy collection from OR, 210. Alpine plants (see Packera) Ant pollination (see Chorizanthe) Apiaceae (see Lomatium) Aquifoliaceae (see //ex) Arizona: Packera franciscana, density and elevational distribution, 213; Peniocereus striatus, plant population and habitat characteristics, 220. Noteworthy collection: Punica granatum, 74. Asparagaceae (see Brodiaea) Asteraceae: Delairea odorata, distribution and communi- ty associations in CA, 85; Ericameria fasciculata, effects of fire on germination, 77; gas exchange rates of savanna sub-shrubs of central TX, 170; Packera franciscana, density and elevational distribution, 213; Pyrrocoma_ liatriformis and P. scaberula, species boundaries from AFLP data, 95. Noteworthy collections: Gutierrezia MEXICO, 211. Astragalus jaegerianus, effects of drought and host plant canopy on, 120. Atriplex hymenelytra, noteworthy collection from MEX- ICO; 211. ramulosa from Barbour, Michael G., dedication of Vol. 57 to, 281 Boraginaceae (see Howellanthus and Plagiobothrys) Brassica oleracea, noteworthy collection from OR, 274. Brassicaceae (see Brassica and Matthiola) Brodiaea matsonii, new sp. from Shasta Co., CA, 261. Cactaceae (see Peniocereus) California: Astragalus jaegerianus, effects of drought and host plant canopy on, 120; Chorizanthe parryi var. fernandina reproductive success, 161; Delairea odorata, distribution and community associations, 85; Dudleya multicaulis, pollination and reproduction in natural and mitigation populations, 42; Ericameria fasciculata, effects of fire on germination, 77; exotic plant invasion in Sequoia sempervirens forests, 11; Plagiobothrys hystriculus, rediscovery, 242; Prosartes parvifolia, tax- onomic status, 129; Sequoia sempervirens stand devel- opment on a 127-year old chronosequence of naturally regenerating forests, 229; Toxicoscordion exaltatum chromosome number, 180; Zeltnera muhlenbergii iden- tity and nomenclature, 184. New taxa: Abies magnifica var. critchfieldii, 141; Brodiaea matsonii, 261; Chenopodium littoreum, 64: Howellanthus dalesianus, 268. Noteworthy collections: Acacia dealbata, 274: Viola howellii, 209. Calyptridium parryi, taxonomy, 145. _ Campanulaceae (see Downingia) Centaurium (see Zeltnera) Chenopodiaceae (see Atriplex and Chenopodium) Chenopodium littoreum, new species from dunes of south- central coastal California, 64. Chorizanthe parryi var. fernandina reproductive success, 161. Chromosome counts: Frasera umpquaensis and F. fasti- giata, 109; Toxicoscordion exaltatum, 180. Compositae (see Asteraceae) Crassulaceae (see Dudleya and Sedum) Cryptic species (see Mentzelia) Cyperaceae (see Schoenoplectus) Delairea odorata, distribution and community associa- tions in CA, 85. Distichlis bajaensis, new species from Baja California, Mexico, 54. Dodonaea viscosa var. angustifolia, noteworthy collection from MEXICO, 211. Downingia: Morphologically cryptic species within D. yina, 20. Dudleya_ multicaulis, pollination and reproduction in natural and mitigation populations, 42. Editors’ Report for Vol. 57, 277. Ericameria fasciculata, effects of fire on germination, 77. Fabaceae: Astragalus jaegerianus, effects of drought and host plant canopy on, 120. Noteworthy collections: Acacia dealbata, from CA, 274; Ornithopus perpusillus, from OR, 275; Prosopis glandulosa var. torreyana, from MEXICO, 211; Trifolium retusum, 275. Fire, effects on germination of Ericameria fasciculata, 77. Frasera umpquaensis and F. fastigiata, differentiation, 106. Gas exchange rates of savanna sub-shrubs of central TX, 170. Gentianaceae (see Frasera and Zeltnera) Gramineae (see Poaceae) Gutierrezia ramulosa, noteworthy collection from MEX- CO, 2 Howellanthus dalesianus, new genus and species, 268. Idaho (see Pyrrocoma and Sedum) Ilex, invasive species and their dispersers in the Pacific Northwest, |. Invasive plants: Delairea odorata, distribution and community associations in CA, 85; J//lex, invasive species and their dispersers in the Pacific Northwest, 1; plant invasion in redwood forest gaps, 11. Isozymes (see Frasera) Keys: Brodiaea spp. with perianth narrowed above the ovary, 266; Calyptridium parryi complex, 158; Down- ingia yina complex, 34; Ilex species growing outside cultivation in Pacific Northwest, 4; Lomatium similar to 280 L. tamanitchii, 207; Mentzelia sect. Bartonia, 259: Plagiobothrys spp. with both prickles and _ bristles, 243: Prosartes of northwest CA and southwest OR, 131: Sedum of ID, 139. Leguminosae (see Fabaceae) Liliaceae (see A//ium and Prosartes) Loasaceae (see Mentzelia) Lomatium tamanitchii, new sp. from OR and WA, 203. Lythraceae (see Punica) Matthioloa incana, noteworthy collection from OR, 274. Melanthiaceae (see Toxicoscordion) Mentzelia sect. Bartonia, new taxa from western No. Am., 246. New taxa: Mentzelia filifolia, M. holmgreniorum, M. lagarosa, M. longiloba var. chihuahuaensis, M. I. var. pinacatensis, M. 1. var. yavapaiensis, M. paradoxensis, M. procera, M. sivinskii, 246. MEXICO: Baja California Norte: New taxon: Distichlis bajaensis, 54. Noteworthy collections: Atriplex hymenelytra, Dodo- naea viscosa var. angustifolia, Gutierrezia ramulosa, Prosopis glandulosa var. torreyana, Tamarix ramo- sissima, 211. Mojave Desert (see Astragalus) Montiaceae (see Calyptridium) Nurse plant (see Peniocereus) Oregon: New taxon: Lomatium tamanitchii, 203. Noteworthy collections: Al/ium peninsulare, 210; Bras- sica oleracea, Matthiola incana, 274; Ornithopus perpusillus, 275; Schoenoplectus californicus, 274; Trifolium retusum, 275. Organ Pipe Cactus National Peniocereus) Ornithopus perpusillus, noteworthy collection from OR, 21D: Monument, AZ (see Packera franciscana, density and elevational distribution, 213: Peniocereus striatus, plant population and habitat char- acteristics, 220. Plagiobothrys hystriculus, rediscovery, 242. MADRONO [Vol. 57 Pinaceae (see Abies) Poaceae (see Distichlis) Polygonaceae (see Chorizanthe) Portulaceae (see Montiaceae) President’s Report for Vol. 57, 276. Prosartes parvifolia, taxonomic status, 129. Prosopis glandulosa var. torreyana, noteworthy collection from MEXICO, 211. Punica granatum, noteworthy collection from AZ, 74. Pyrrocoma liatriformis and P. scaberula, species bound- aries from AFLP data, 95. Redwood forest (see Sequoia) Reviews: Desert Wisdom/Agaves and Cacti: CO>, Water, Climate Change by Park S. Nobel, 73. Salmon River Cyn, ID (see Sedum) Sapindaceae (see Dodonaea) Savannas: Gas exchange rates of three sub-shrubs of central TX, 170. Schoenoplectus californicus, noteworthy collection from OR, 274. Sedum valens, new sp. from Salmon River Cyn, ID, 136. Sequoia sempervirens, exotic plant invasion in redwood forest gaps, 11; stand development on a 127-year old chronosequence of naturally regenerating forests, 229. Sierra Nevada, CA (see Abies) Siskiyou Mts (see Prosartes) Tamaricaceae (see Tamarix) Tamarix ramosissima, noteworthy collection from MEX- [CO. 2145 Taxodiaceae (see Sequoia) Texas, savanna sub-shrub gas exchange rates, 170. Trifolium retusum, noteworthy collection from OR, 275. Toxicoscordion exaltatum chromosome number, 180. Vernal pools (see Plagiobothrys) Viola howellii, noteworthy collection from CA, 209. Washington (see Lomatium and Pyrrocoma) Zeltnera muhlenbergii identity and nomenclature, 184. MADRONO, Vol. 57, No. 4, pp. 281—282, 2010 DEDICATION MICHAEL G. BARBOUR The California Botanical Society dedicates this volume of Madrono to one of California’s most influential plant ecologists, Michael G. Barbour, in recognition of his distinguish accomplishments in research, in teaching, and in conservation. Michael was born in February, 1942. After graduating magna cum laude with a BS in Botany in 1963 from Michigan State University, Barbour went on to obtain his doctorate in Botany from Duke University in 1967. Prior to obtaining his doctorate, he was also a Fulbright Fellow in Adelade, Australia. Michael joined the faculty of the University of Califor- nia, Davis (UCD) in 1967 and became one of the youngest tenured faculty at UCD. During his 40 years at UCD, he was a professor in a succession of departments (Botany, Plant Biology, Ornamental Horticulture, and finally Plant Sciences, from which position he retired in 2008. Although California has been his home throughout his career, and has provided the mainstay of his academic pursuits, Michael has conducted or supervised research in many parts of the world, including Argentina, Spain, Portugal, Israel, Mexico, and Australia. A common thread in his career has been exploring the stressors, disturbances, and tolerances that define the habitat limits of different species and vegetation communities. His work on desert scrub in the southwestern deserts of North America investigated the role of competition for resources in the spatial patterning of species as well as the biogeographic and genetic relationships between Larrea in North and South America. His work on salt tolerance of Californian coastal dune and salt marsh plants underscored the interaction of physiology and tolerance in the zonal distribution of species and communities. His work in the red and white fir forests of the Sierra Nevada of California investigated the means by which dominant species in cold and snowy environ- ments displace each other in the major Sierran ecotone at 2000 m elevation. Working in the Canary Islands, he studied the age structure, stand dynamics, and fire regime of old-growth Pinus canariensis forests—a species with a unique set of attributes for a serotinous conifer. Most recently, he completed a study of mixed evergreen forests in central Spain and northern California, teasing out the environ- mental factors that seemed to explain varying dominance by conifers vs. hardwoods and by particular species, such as Quercus pyrenaica and Q. garryana. Research results have been published in the American Journal of Botany, American Midland Naturalist, the Journal of Biogeography, Forest Ecology and Manage- ment, Madrono, the Journal of Vegetation Science, Conservation Biology, Israel Journal of Botany, Oeco- logia, and as chapters in several technical monographs. Michael has published more than 150 books, chapters, _ and papers. | Michael has taken his academic role seriously beyond _ his own research. As an educator he has authored or co- _ authored a diversity of text books on botany, plant _ ecology, landscape ecology, and vegetation. He took on the Vegetation of North America with his major professor at Duke, W.D. Billings, and has also co- authored the most widely used texts on plant ecology. He and his long-time UCD friend and mentor, Jack Major, edited and wrote portions of the first two editions of the widely used compendium on vegetation ecology for California (the Terrestrial Vegetation of California), which set the stage for many developments in the study of California and western North American vegetation. Michael is also a truly gifted teacher. His courses on California plant communities and plant ecology are legendary, involving an expertly crafted marriage of field trips and lectures. His lectures are pitch-perfect matches with his audience, whether laypeople hearing about vernal pools for the first time, or graduate students, learning about the fine points of fire ecology or physiological ecology. Watching him teach, providing clear summary points and artistic real-time chalk board illustrations (he is not too fond of PowerPoint), has been for me a humbling lesson in the art of classical instruction. His capabilities did not go unrecognized by UCD. In 1988, the campus awarded him a Citation for Distinguished Teaching. Michael served as the major professor for 50 graduate students and was a committee member for many others. A number of his ex-students now have important faculty, NGO, federal and state agency positions, or are successful consultants. Though eternally congenial, even-handed, and humor- ous, Michael has not shied away from academic debate or conflict. When challenged by European colleagues to get American colleagues to use European methods of vegetation analysis and classification after supporting the development of a new classification system for Califor- nia’s vegetation he invited them over to work with him. As a result, he involved many phytosociologists in the last two decades in the rapid collaborative accumulation of knowledge on our state’s vegetation classification and has continued to be a voice for philosophical exchange in Photo by Brett Hall, September 2009. 282 the International Association for Vegetation Science. His spirit of information-sharing and openness prevailed in many forms throughout Michael’s career. It was important, for example, in his role as the first chair of the Ecological Society of America’s Vegetation Panel, where the task was to develop and standardize a single classification system that would be scientifically ground- ed and adopted by all federal natural resource agencies. Michael, though academically trained and accom- plished, also became involved in a number of conserva- tion issues and has shown many of us the power of promoting conservation through scientifically defensible means. My first encounter with this was as a member of the California Native Plant Society’s Plant Communities Committee where he deftly and calmly (though he refutes this) led a disparate group of opinionated ecologists to a shared vision of conservation-based, quantitative vege- tation classification. He similarly became involved in an effort to develop a quantifiable approach to vernal pool classification monitoring and evaluation, and from that to putting metrics on vernal pool restoration by quantifying the deviance between created vernal pool communities and naturally occurring ones. After a MADRONO [Vol. 57 decade of work with a team of vernal pool experts, his approach has now been adopted by the U.S. Environ- mental Protection Agency and other regulatory agencies. Dr. Barbour is a lucid writer and a perceptive editor, having served on the editorial board of several peer- reviewed journals. He is also not above writing and collaborating on projects that are more literary than they are scholarly. He has co-authored and co-performed with poet Gary Snyder and has co-written several popular articles with his wife Valerie Whitworth on California conservation and natural history. In short, Michael is worthy of all the academic praise he has received, but equally as worthy in his role as a bridger of schisms, a practical, yet passionate spokesperson for the marvelous beauty of nature, and as an all-around citizen of the world. Michael, it is a pleasure to be given the honor to write this dedication to you. Todd Keeler-Wolf, Ph.D. Senior Ecologist Vegetation Classification and Mapping Program California Department of Fish and Game MADRONO A WEST AMERICAN JOURNAL OF BOTANY VOLUME LVII 2010 BOARD OF EDITORS Class of: 2010—FRED HRUSA, California Department of Food and Agriculture, Sacramento, CA RICHARD OLMSTEAD, University of Washington, Seattle, WA 2011—JAMIE KNEITEL, California State University, Sacramento, CA KEVIN RICE, University of California, Davis, CA 2012—-GRETCHEN LEBUHN, San Francisco State University, CA ROBERT PATTERSON, San Francisco State University, CA 2013—-ERIC ROALSON, Washington State University, WA KRISTINA SCHIERENBECK, California State University, Chico, CA Corresponding Editor—TIMOTHY LOWREY Museum of Southwestern Biology MSC03 2020 University of New Mexico Albuquerque, NM 87131-0001 madrono@unm.edu AND Copy Editor—RICHARD WHITKUS Department of Biology Sonoma State University 1801 E. Cotati Avenue Rohnert Park, CA 94928-3609 whitkus@sonoma.edu Published quarterly by the California Botanical Society, Inc. Life Sciences Building, University of California, Berkeley 94720 Printed by Allen Press, Inc., Lawrence, KS 66044 MADRONO VOLUME 57 TABLE OF CONTENTS Allen, Robert L. (see Jones, C. Eugene, Frances M. Shropshire, Robert L. Allen and Youssef C. Atallah) Allen, Robert L. (see also Jones, C. Eugene, et al.) Anderson, Greta, Sue Rutman and Seth M. Munson, Plant population and habitat characteristics of the endemic Sonoran Desert cactus Peniocereus striatus in Organ Pipe Cactus National Monument, PTZ OU 2 re Sc Saeco ee pede a Poe Rn oh ee Atallah, Youssef C. (see Jones, C. Eugene, Frances M. Shropshire, Robert L. Allen and Youssef C. Atallah) Atallah, Youssef C. (see also Jones, C. Eugene, et al.) Bell, Hester L., A new species of Distichlis (Poaceae, Chloridoideae) from Baja California, Mexico Bencie, Robin (see Mesler, Michael) Benet-Pierce, Nuri, and Michael C. Simpson, Chenopodium littoreum (Chenopodiaceae), a new goosefoot from dunes of south-central coastal California nn Bjork, Curtis R., Sedum valens (Crassulaceae), a new species from the Salmon River Canyon of Idaho ____ Bjork, Curtis R. (see also Smith, James F.) Blair, Brent C., Deborah K. Letourneau, Sara G. Bothwell and Grey F. Hayes, Disturbance, resources, and exotic plant invasion: gap size effects in a redwood forest Bothwell, Sara G. (see Blair, Brent C.) Brainerd, Richard E. (see Darrach, Mark, et al.) Burk, Jack H. (see Jones, C. Eugene, et al.) Darrach, Mark, et al., Lomatium tamanitchii (Apiaceae) a new species from Oregon and Washington state, USA Denslow, Michael W., Gabrielle Katz and Wendy Hodgson, Noteworthy collection from Arizona ________. Detka, Jon R., and Susan C. Lambrecht, Effects of fire on germination of Ericameria fasciculata (Asteraceae), a rare maritime chaparral shrub - eee es i eee hee eee ett ea en DiTomaso, Joseph M. (see Robison, Ramona) Fowler, James F., and Carolyn Hull Sieg, Density and elevational distribution of the San Francisco Peaks ragwort, Packera franciscana (Asteraceae), a threatened single-mountain endemic Furuya, Mitsuru, and O.W. Van Auken, Gas exchange rates of three sub-shrubs of central Texas savannas __. (sarcia, Peter J. Noteworthy collections from: Mexico -.2..3.2..05.) 2 eee a ee Glenne, Gina (see Smith, James F.) Guilliams, C. Matt (see Simpson, Michael G., Michael Silveira and C. Matt Guilliams) Halse, Richard R., Noteworthy collections from Oregon _ Hayashi, Bianca (see Mesler, Michael) Hayes, Grey F. (see Blair, Brent C.) Hipkins, Valerie (see Wilson, Barbara L.) Hodgson, Wendy (see Denslow, Michael W.) Hufford, Larry (see Schenk, John J.,) Huggins, T. R., B. A. Prigge, M. R. Sharifi and P. W. Rundel, The effects of long-term drought on host plant canopy condition and survival of the endangered Astragalus jaegerianus (Fabaceae) Jones, C. Eugene, Frances M. Shropshire, Robert L. Allen and Youssef C. Atallah, Pollination and reproduction in natural and mitigation populations of the many-stemmed dudleya, Dudleya multicaulis (Crassulaceae) Jones, C. Eugene, et al., Do native ants play a significant role in the reproductive success of the rare San Fernando Valley spineflower, Chorizanthe parryi var. Fernandina (Polygonaceae) ___------...-... Katz, Gabrielle (see Denslow, Michael W..,) Kaye, Tom N. (see Wilson, Barbara L.) Keeler-Wolf, Todd, Dedication of Volume 57 to Michael G. Barbour Lambrecht, Susan C. (see Detka, Jon R.) Lanner, Ronald M., Abies magnifica var. critchfieldii, a new California red fir variety from the Sierra Nevada ___ Letourneau, Deborah K. (see Blair, Brent C.) Lindstrand, Len, HI, Noteworthy collection from California = gett eea ie noe See Little, “R; John, Noteworthy collechom from California. ..2:2.3.. 2. needed e eee ee Longstreth, David J., Review of Desert Agaves and Cacti: COs Water, Climate ‘Change by Park S. Nobel Lowry, Tim, and Richard Whitkus, Editors’ Report for Volume 57 - Luttrell, Jim (see Jones, C. Eugene, et al.) McNeal, Dale W., and Wendy B. Zomlefer, Documentation of the chromosome number for the California endemic, Toxicoscordion exaltatum (Liliales: Melanthiaceae) Mesler, Michael, Robin Bencie and Bianca Hayashi, A resurrection for Siskiyou bells, Prosartes parvifolia (Liliaceae), a rare Siskiyou Mountains endemic sss ey nee ree Peer ee Michels, Kristin Hageseth (see Russell, Will) Mousseaux, Mark, Noteworthy collection from Oregon ____ ao £2 eee nee eee Munson, Seth M. (see Anderson, Greta) 220 54 64 136 11 203 74 515) 213 170 211 274 120 42 { 2010] TABLE OF CONTENTS Otting, Nick (see Darrach, Mark, et al.) Parker, V. Thomas, President’s Report for Volume 57 Patterson, Robert (see Walden, Genevieve K.) Perkins, Dusty N. (see Smith, James F.) Preston, Robert L., Brodiaea matsonii, (Asparagaceae: Brodiaeoideae) a new species from Shasta County, California ___. et ee Preston, Robert L., Brad D. Schafer and Margaret Widdowson, Rediscovery of Plagiobothrys hystriculus (Boraginaceae) and notes on its habitat and associates Prigge, B. A. (see Huggins, T. R.) Pringle, James S., The identity and nomenclature of the Pacific North American species Zeltnera muhlenbergii (Gentianaceae) and its distinction from Centarium tenuiflorum and other species with which it has been confused ___ Sas oat : Robison, Ramona, and Joseph M. DiTomaso, Distribution and community associations of Cape ivy (Delairea odorata) in California Rundel, P. W. (see Huggins, T. R.) Russell, Will, and Kristin Hageseth Michels, Stand development on a 127-yr chronosequence of naturally regenerating Sequoia sempervirens (Taxodiaceae) forests Rutman, Sue (see Anderson, Greta) Sandquist, Darrren R. (see Jones, C. Eugene, et al.) Schafer, Brad D. (see Preston, Robert L., Brad D. Schafer and Margaret Widdowson) Schenk, John J., and Larry Hufford, Taxonomic novelties from western North America in Mentzelia section Sharifi, M. R. (see Huggins, T. R.) Shropshire, Frances M. (see Jones, C. Eugene, Frances M. Shropshire, Robert L. Allen and Youssef C. Atallah) Shropshire, Frances M. (see also Jones, C. Eugene, et al.) Sieg, Carolyn Hull (see Fowler, James F.) Silveira, Michael (see Simpson, Michael G., Michael Silveira and C. Matt Guilliams) Simpson, Michael G., Michael Silveira and C. Matt Guilliams, Taxonomy of calyptridium parryi Smith, James F., Dusty N. Perkins, Curtis R. BjOrk and Gina Glenne, Species boundaries in Pyrrocoma liatriformis and Pyrrocoma scaberula (Asteraceae) based on AFLP data Song, Leo C., Jr. (see Jones, C. Eugene, et al.) Thie, Kirsta K. (see Darrach, Mark, et al.) Van Auken, O. W. (see Furuya, Mitsuru) Walden, Genevieve K., and Robert Patterson, Howellanthus dalesianus, recognition of a new genus and species in the tribe Phacelieae (Boraginaceae) Walker, Sean E. (see Jones, C. Eugene, et al.) Whitkus, Richard (see Lowry, Tim) Widdowson, Margaret (see Preston, Robert L., Brad D. Schafer and Margaret Widdowson) Wilson, Barbara L., Valerie Hipkins and Tom N. Kaye, One taxon or two: Are Frasera umpquaensis and F- fastigiata (Gentianaceae) distinct species? Wilson, Barbara L. (see also Darrach, Mark, et al.) Zika, Peter F., Invasive hollies (/ex, Aquifoliaceae) and their dispersers in the Pacific Northwest Zomlefer, Wendy B. (see McNeal, Dale W.) DATES OF PUBLICATION OF MADRONO, VOLUME 57 Number 1, pages 1—76, published 31 August 2010 Number 2, pages 77-144, published 30 September 2010 Number 3, pages 145-212, published 21 December 2010 Number 4, pages 213-282, published 4 May 2011 ill 246 20 145 oe ot _ 4 = heey SUBSCRIPTIONS — MEMBERSHIP The California Botanical Society has several membership types (individuals ($35 per year; family $40 per year; emeritus $27 per year; students $27 per year for a maximum of 7 years). Late fees may be assessed. Beginning in 2011, rates will increase by $5 for all membership types except life memberships, for which rates will increase by $100, and student memberships, which will not show a rate increase. Members of the Society receive MADRONO free. Institutional subscriptions to MADRONO are available ($70). Membership is based on a calendar year only. Life memberships are $750. Applications for membership (including dues), orders for subscriptions, and renewal payments should be sent to the Membership Chair. Requests and rates for back issues, changes of address, and undelivered copies of MADRONO should be sent to the Corresponding Secretary. INFORMATION FOR CONTRIBUTORS Manuscripts submitted for publication in MADRONO should be sent to the editor preferably as Microsoft Word (.doc), Rich Text Format (.rtf), or Portable Document Format (.pdf) files. It is preferred that all authors be members of the California Botanical Society. Manuscripts by authors having outstanding page charges will not be sent for review. Manuscripts may be submitted in English or Spanish. English-language manuscripts dealing with taxa or topics of Latin America and Spanish-language manuscripts must have a Spanish RESUMEN and an English ABSTRACT. For all articles and short items (NOTES, NOTEWORTHY COLLECTIONS, POINTS OF VIEW, etc.), fol- low the format used in recent issues for the type of item submitted. Allow ample margins all around. Manuscripts MUST BE DOUBLE-SPACED THROUGHOUT. For articles this includes title (all caps, centered), author names (all caps, centered), addresses (caps and lower case, centered), abstract and resumen, five key words or phrases, text, acknowledgments, literature cited, tables (caption on same page), and figure captions (grouped as consecutive paragraphs on one page). Order parts in the sequence listed, ending with review copies of illustrations. The title page should have a running header that includes the name(s) of the author(s), and a shortened title. Avoid foot- notes except to indicate address changes. Abbreviations should be used sparingly and only standard abbreviations will be accepted. Table and figure captions should contain all information relevant to information presented. All measurements and elevations should be in metric units, except specimen citations, which may include English or metric measurements. Authors are encouraged to include the names, addresses, and e-mail addresses of two to four potential reviewers with their submitted manuscript. Authors of accepted papers are required to submit an electronic version of the manuscript. Microsoft Word 2000 or later or WordPerfect 9.0 (or later) for Windows is the preferred software. Line copy illustrations should be clean and legible, proportioned to the MADRONO page. Scales should be in- cluded in figures, as should explanation of symbols, including graph coordinates. Symbols smaller than | mm after reduction are not acceptable. Maps must include a scale and latitude and longitude or UTM references. Presentation of nomenclatural matter (accepted names, synonyms, typification) should follow the format used by Sivinski, Robert C., in MADRONO 41(4), 1994. Institutional abbreviations in specimen citations should follow Index Herbariorum. 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 California Botanical Society members current in the volume year that their contributions are published are allowed five free pages per volume year. Additional pages will be charged at the rate of $40 per page. Joint authors may apply all or a portion of their respective five-page allotments to a jointly-published article. Partial pages will be charged as full. The purpose of this fee is not to pay directly for the costs of publishing any particular paper, but rather to allow the Society to continue publishing MADRONO on a reasonable schedule, with equity among all members for access to its pages. Printer’s fees for color plates and other complex matter (including illustrations, charts, maps, photographs) will be charged at cost. Author’s changes after typesetting @ $4.50 per line will be charged to authors. Page charges are important in maintaining Madrono as a viable publication, and timely payment of charges 1s appreciated. At the time of submission, authors must provide information describing the extent to which data in the manuscript have been used in other papers that are published, in press, submitted, or soon to be submitted elsewhere. 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