VOLUME 59, NUMBER | JANUARY-MARCH 2012 MADRONO A WEST AMERICAN JOURNAL OF BOTANY POINTS OF VIEW COMMENT ON THE GABBRO SOILS OF PINE HILL | ON AS oy WN 08) 218 A Re Cee eee Eee ee ree TONy A ee ne ere | CONTENTS VASCULAR ALPINE FLORA OF MOUNT WASHBURN, YELLOWSTONE NATIONAL Park, USA Ken Aho and Janet Bala vrcccccccccccecccccnceciscado¥Ouod ve seson os vga phe BRR coscesesnvesvos 2 IMPACT OF RECURRENT FIRE ON ANNUAL PLANTS: A CASE STUDY FROM THE WESTERN EDGE OF THE COLORADO DESERT Robert J. Steers and Edith B: Alert > 0.0.6. 2365 Ee voc nccccaceccccccesecces 14 STATUS OF BINGHAM’S MORNING-GLORY IN THE LIGHT OF ITS REDISCOVERY R. K. Brummitt, Scott D. White, and Justin M. WOO ..........cccccceceueceneceues 25 CHANGE IN RANK OF ERIODICTYON TRASKIAE SUBSP. SMITHII (HYDROPHYLLACEAE) GOCE AEANTON EN TIAN, oo Li CAS ees hanson NUCL NE oRiae IEG cco ee asseeseeeees 28 NEW SPECIES MIMULUS SOOKENSIS (PHRYMACEAE), A NEW ALLOTETRAPLOID SPECIES DERIVED FROM MIMULUS GUTTATUS AND MIMULUS NASUTUS Beverly G. Benedict, Jennifer L. Modliszewski, Andrea L. Sweigart, Noland H. Martin, Fred R. Ganders, and John H. WiIIIiS ..... 000.00. ccecceeeee 29 BOOK REVIEWS NORTHWEST CALIFORNIA: A NATURAL HISTORY TO) PUI CH rea S TEI cis Se cd teers TID otto Sgn I aoa t ache ee acne eed 44 PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICA OIE? MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and 1s issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. POSTMASTER: Send address changes to MADRONO, Kim Kersh, Membership Chair, Uni- versity and Jepson Herbarium, University of California, Berkeley, CA 94720-2465. kersh @berkeley.edu. Corresponding Editor—MatTT RITTER Copy Editor—RICHARD WHITKUS Biological Sciences Department Department of Biology Cal Poly, San Luis Obispo Sonoma State University 1 Grand Avenue 1801 E. Cotati Avenue San Luis Obispo, CA 93407 Rohnert Park, CA 94928-3609 madronoeditor @ gmail.com whitkus @sonoma.edu Book Editor—Mat?T RITTER Noteworthy Collections Editors—DI&TER WILKEN Board of Editors Class of: 2012—GRETCHEN LEBUMN, San Francisco State University, CA ROBERT PATTERSON, San Francisco State University, CA 2013—Eric ROALSON, Washington State University, WA KRISTINA SCHIERENBECK, California State University, Chico, CA 2014—BRANDON PRATT, California State University, Bakersfield, CA Tom WENDT, University of Texas, Austin, TX CALIFORNIA BOTANICAL SOCIETY, INC. OFFICERS FOR 201 1—2012 President: V. Thomas Parker, Department of Biology, San Francisco State University, San Francisco, CA 94132, parker @sfsu.edu First Vice President: Andrew Doran, University and Jepson Herbaria, University of California, Berkeley, CA 94720, andrewdoran @ berkeley.edu Second Vice President: Vacant. Recording Secretary: Michael Vasey, Department of Biology, San Francisco State University, San Francisco, CA 94132, mvasey @sfsu.edu Corresponding Secretary: Anna Larsen, Jepson Herbarium, University of California, Berkeley, CA 94720, secretary @calbotsoc.org Treasurer: Thomas Schweich, California Botanical Society, Jepson Herbarium, University of California, Berkeley, CA 94720, tomas @schweich.com The Council of the California Botanical Society comprises the officers listed above plus the immediate Past President, Dean Kelch, Jepson Herbarium, University of California, Berkeley, CA 94720, dkelch@berkeley.edu; the Membership Chair, Kim Kersh, University and Jepson Herbaria, University of California, Berkeley, CA 94720, kersh @berkeley.edu; the Editor of Madrono; and three elected Council Members: Chelsea Specht, Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720-2465, cdspecht@berkeley.edu; Ellen Simms, Department of Intergrative Biology, 1005 Valley Life Sciences Bldg., #3140, University of California, Berkeley, CA 94720, esimms @berkeley.edu. Staci Markos, University and Jepson Herbaria, University of California, Berkeley, CA 94720, smarkos @berkeley.edu. Graduate Student Representatives: Genevieve Walden, Department of Integrative Biology and Jepson Herbarium, University of California, Berkeley, CA 94720, gkwalden@ gmail.com. Administrator: Lynn Yamashita, University of California, Berkeley, CA 94720, admin @calbotsoc.org. Webmaster: Ekaphan (Bier) Kraichak, University of California, Berkeley, CA 94720, ekraichak @ gmail.com. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). MADRONO, Vol. 59, No. 1, p. 1, 2012 POINT-OF-VIEW COMMENT ON THE GABBRO SOILS OF PINE HILL Burge and Manos (2011) investigated the genetic relationships of Ceanothus roderickii W. Knight and Ceanothus cuneatus Nutt. var cunea- tus and sampled surface soils where the plants were found. They claimed to have shown that the two species are associated with chemically different gabbro soils. The mineralogical and chemical differences among gabbro rocks are great and plant distri- butions from those dominated by olivine to those dominated by Ca-feldspars might be expected to be different. With respect to C. roderickii and C. cuneatus var. cuneatus, there are three key questions: (1) What is the range of gabbro soils on which C. roderickii will grow? (2) What is the range of gabbro soils on which C. cuneatus var. cuneatus will grow? (3) Considering gabbro rocks and soils where the ranges of C. roderickii and C. cuneatus var. cuneatus do not overlap, what are the mineralogical differences in the rocks and the chemical differences in the soils that might limit plant distributions. A final test would be to plant the two species in soils from different kinds of gabbro rocks under climatically similar or controlled. Burge and Manos (2011) did not identify the specific gabbro rock mineralogies; they sampled only surface soils, and they ascertained only the readily extractable portions of the chemical elements. Locations where they sampled surface soils may have been in areas where the distribu- tions of C. roderickii and C. cuneatus var. cuneatus do not overlap, but the methods were not adequate to distinguish different kinds of rocks and soils. Their data indicate that the greatest differences between the surface soils at sites with different ceanothus species were differ- ent amounts of Mehlich III (dilute acids and EDTA) extractable P. Alexander (2011) sampled the parent rocks and both surface (O—15 cm) and subsoils (30—45 cm) at one site with C. roderickii and two sites without it on the Pine Hill gabbro. Phosphorus was ascertained from aqua regia digestion of the soils to evaluate the total elemental reserves in the soils. The soil with C. roderickii had subsoil P similar to that in the other soils, but the surface soil in the C. roderickii plant community had much more P than in the surface soils at the sites lacking C. roderickii. The surface soil at the C. roderickii site also had much more organic matter than the soils at the other two sites. Evidently, the amounts of P in the surface soils was largely dependent on the amounts of plant detritus that had been incorporated into them, which is a function of entire ecosystems, not only a single species. Perhaps the soil parent materials at the C. roderickii sites where Burge and Manos sampled the surface soils had as much P as the parent materials of other gabbro soils, but the plant communities at C. roderickii sites were cycling less P than the plant communities at the wedgeleaf ceanothus sites? Unfortunately, the methods of Alexander (2011) are too intensive to apply broadly and the low- intensive methods of Burge and Manos are inadequate to show gabbro petrologic and soil differences related to the distributions of endemic plants. Perhaps future investigations that are less intensive than that of Alexander, but comprehensive enough to identify the kinds of gabbro parent rocks and both surface and subsoil reserves of key elements, will identify what gabbro rock and soil features lead to different plant distributions. —EARL B. ALEXANDER, Soils and Geoecol- ogy, 1714 Kasba Street, Concord, CA 94518; alexandereb@att.net. LITERATURE CITED ALEXANDER, E. B. 2011. Gabbro soils and plant distributions on them. Madrono 58:113-—122. BURGE, D. O. AND P. S. MANOos. 2011. Edaphic ecology and genetics of the gabbro-endemic shrub Ceanothus roderickii (Rhamnaceae). Madrono 58:1-21. MADRONO, Vol. 59, No. 1, pp. 2-13, 2012 VASCULAR ALPINE FLORA OF MOUNT WASHBURN, YELLOWSTONE NATIONAL PARK, USA KEN AHO! Department of Biology, Montana State University, Bozeman, MT 59717 ahoken@isu.edu JANET BALA Idaho State Herbarium (IDS), Idaho State University, Pocatello, ID 83209 ABSTRACT Mount Washburn, the principal peak in the volcanic Washburn Range, is an important site for both tourism and research in Yellowstone National Park. This paper provides: 1) descriptions of plant community types on Mt. Washburn, 2) biogeographic comparisons of species diversity for several ranges in the North-Central Rockies, and 3) an annotated species list of the alpine vascular flora, including summaries of constancy, local abundance, and preferred habitats. The alpine flora consists of one hundred and twenty-six vascular plant species from seventy-five genera and twenty-eight families. Biogeographic analyses suggest that the flora is depauperate for the region, with relatively low rates of colonization. These results agree with the predictions of the theory of island biogeography for small isolated ecosystems, and emphasize the vulnerability of Washburn to sub-alpine encroachment as the result of climate change. Key Words: Alpine flora, andesitic substrates, biogeography, Mount Washburn, Yellowstone National Park. Mount Washburn (3124 m), a volcanic for- mation in north-central Yellowstone National Park (YNP), has long been an important des- tination for tourism and _ scientific research. Washburn is one of the most frequently climbed alpine summits in the Rocky Mountains (Aho and Weaver 2010). Previous scientific research on Mt. Washburn includes studies of geology (Feeley et al. 2002), conifer distributions (Kokaly et al. 2003), whitebark pine ecology (Weaver and Dale 1974; Mattson and Reinhart 1990; Tomback et al. 2001), and grizzly bear ecology (Podruzny 1999). While alpine vegetation has been described for volcanic substrates in the coastal Cordillera (Douglas and Bliss 1977; Hunter and Johnson 1983) and southern Rocky Mountain regions (Baker 1983; Rottman and Hartman 1985; Taye 1995; Seagrist and Taylor 1998), comparable des- criptions for northern Rocky Mountain volcanic peaks are scarce. Aho and Weaver (2010) identified distinct alpine communities on Mt. Washburn, and described community evolutionary trends. This work, however, provided neither a formal inven- tory of Washburn alpine species, nor a comparison of the Washburn flora to those of other alpine locations. Annotated species lists are valuable tools for monitoring/management (O’Kane 1988), hy- pothesis generation (Bell and Johnson 1980), ‘Present address: Department of Biology, Idaho State University, Pocatello, ID 83209. and floristic comparisons (Baker 1983). The absence of an inventory for Mt. Washburn is notable given the existence of such lists for the Beartooth Mountains to the north (Johnson and Billings 1962; Lackschewitz 1994), the Tetons to the south (Spence and Shaw 1981), and the Madison, Gallatin, and Tobacco Root Moun- tains to the west (Pemble 1965; Cooper et al. 1997). The Mt. Washburn alpine zone may have been overlooked because of its insular characteristics (i.e., small size and isolation; cf. Billings 1978). The extent of Mt. Washburn alpine vegetation is less than 1.2 km? (Despain 1990), while the nearest neighboring areas of alpine vegetation are in the region of Thunderer Peak, approximately 30 km to the northeast (Fig. 1). The insularity of the Washburn alpine is notable since it may result in increased vulnerability to subalpine encroach- ment as a result of climate change (cf. Hadley 1987; Bruun and Moen 2003; Halloy and Mark 2003). This paper describes the flora of the alpine zone of Mt Washburn (not subalpine zones, nor the more general Washburn Range). First, it describes the alpine communities and environ- ments of Mt. Washburn, including comparisons to other alpine ranges, particularly those on andesitic substrates. Second, biogeographic anal- yses of species diversity are presented to provide a regional context for the Washburn alpine flora. Third, an annotated alpine vascular species list is provided, based on both current and historical collections. 110°27'30"W 110°27'°0"W 110°26'30"W 760 0 190 380 _— Kilometers 110°0'9"'W / Fisher Cr. : Beartooth Lk. 0 20 40 Kilometers bn 110°0'0"W 111°0'0"W Ht 7 iA NORA Raw R ‘ Why te ®R\\\ \ \ \ Mt i | ANN NY vy} }Y \ \ NV EM NY WAY’ 110°30'0"W AHO AND BALA: ALPINE FLORA OF MOUNT WASHBURN 3 110°26'0"W 110°25'30"W 110°25'0"W. 110°24'30"W 110°24'0"W 110°23'30"W ee [7 SN : NAAN \ a cs ii / . ‘ LN fA \ \\ -— > aM 4 TAN WU AA Le ie Le Tk eee (NAL s Sy} 1 eS i < °49'0" Sh AA /Surveyed/Ardas 477" Sen Za a Vii Dai a So PRA wy dn J LEN PYG ER a sea Wi 110°0'0"W Thunderer Peak Mount Washburn \ { Parker Peak . Canyon Village YELLOWSTONE NATIONAL PARK 0255. °10 35 20 a ee Kilometers 111°0'0O"W Fic. 1. 110°30'0"W 110°0'0"W Maps of study area. (a) Overview of Yellowstone National Park (YNP) with respect to Montana, Wyoming, and Idaho. (b) View of YNP showing the Washburn study area and three locations for which year-round alpine weather data is available. (c) Close up view of Washburn summit. The extent of Washburn alpine vegetation 1s from a ARC-GIS shapefile based on the vegetation classification of YNP by Despain (1990). (d) Alpine vegetation areas in Yellowstone National Park. METHODS Study Area Mount Washburn (3124 m) is the highest peak in the Washburn Range, a volcanic formation in north-central Yellowstone National Park (44°48'N, 110°26’W; Fig. 1). The area above treeline (>2950 m) is small (1.2 km’), and dominated by cushion plants, perennial forbs, and deep rooted graminoids (Aho and Weaver 2010). The plant-supporting surficial rock of the Washburn Range is from the Langford Forma- tion of the Thorofare Creek Group, a unit of the Absaroka Volcanic Supergroup (Smedes and Protska 1972). The Langford Formation consists of both light colored lava flows and alluvial facies composed of hornblende and pyroxene andesite 4 MADRONO fragments deposited between 47-49 million years ago (Smedes and Prostka 1972). On Mount Wash- burn the Langford Formation unconformably overlies strata of the Washburn Group, the oldest member of the Absaroka Volcanic Supergroup. Glaciers, most recently from the Pinedale Glaci- ation have scoured the Washburn Range resulting in the present-day rounded appearance of its ridges and northern slopes (Pierce 1979). Detailed year-round and seasonal weather for Mt. Washburn is summarized in Aho and Weaver (2010). These data indicate that precipitation on Washburn is lower than at adjacent alpine ranges (i.e., <800 mm yr '). The mean frost-free season length (number of days with min temps >0°C) on Mt. Washburn is 93 days. This is comparable to other nearby alpine and high subalpine sites (Aho 2006). Voucher Collection Vascular alpine species on Mount Washburn were inventoried during growing seasons (approx. June 25—-Aug 30) over 5 years (2000-2004). During this period thirty-four one day collections were made from four contiguous summits that constitute the majority of the Mt. Washburn alpine (Fig. 1). As species were collected, voucher data were recorded, as well as qualitative infor- mation concerning species constancy, local abun- dance, moisture regime, and species association with eight broad habitat types. These habitat types were: 1) dense turf, 2) rocky turf, 3) ridgetops, 4) talus/scree, 5) late melting snow- banks, 6) ledges on south-facing cliffs, 7) dis- turbed, and 8) treeline. Types 1-6 have been previously recognized as distinct nodal commu- nities on Mt. Washburn (Aho and Weaver 2010). Turf, ridgetops, talus, and snowbanks sites are well documented circumboreal alpine ecosystem components which often contain distinct commu- nities (Billings 2000; Korner 2003). The “‘dis- turbed”’ habitat included areas such as roads, trails, and structures which are frequent through- out the Mt. Washburn alpine. The “‘treeline”’ habitat constituted subalpine/alpine ecotonal sites. Following field collection and identification, voucher specimens were deposited at the Yellow- stone National Park herbarium (YELLO) in Gardiner, WY, and at the Idaho State herbarium (IDS) in Pocatello, ID. To provide a comprehensive species list, our inventory includes not only species collected in 2000-2004, but those collected in the alpine zone by others and vouchered at YELLO over the last 90 years (the earliest vouchers from Mt. Wash- burn date from 1922). The quality of voucher labels dictated the degree to which environments for these species could be described. Species whose vouchered location was uncertain are not included [Vol. 59 here. Nomenclature and IDs for all species follows Dorn (2001). Biogeographic Analyses To estimate the effects of alpine size and iso- lation in the region, plots were established on Mt. Washburn (36 plots), and in the alpine of two other ranges: the Northern Absarokas (82 plots; 9 peaks), and the Beartooth Plateau (60 plots; 6 peaks). The three ranges are adjacent (Fig. 1), but differ widely in their planar area above treeline (1.2 km’, 2384 km’, and 768 km? for Washburn, the Northern Absarokas, and the Beartooths respectively; Hadley 1987), and distance to other alpine ranges (>50 km for Washburn, and <5 km for the other two ranges; Aho 2006). Each plot consisted of ten 20 < 50 cm subplots situated at each meter on a 10 m line. In each subplot ocular estimates of cover were made for each vascular species. Plots were established randomly within each of five environments (N face, S face, ridgetop, talus, and late-melt). Whenever possible the en- vironments on each mountain range were sampled in the same proportions, 1.e., each environment made up approximately 20% of total number of samples from each range. To compare richness of floras, species area curves were constructed from these data using first order jackknife procedures (Palmer 1990). To compare the importance of rare species, rank abundance dominance (RAD) plots were fit with Preston log-normal models (Preston 1948), this approach often effectively describes local com- munity rank/dominance patterns (Hubbell 2001). Jackknife and RAD analyses were conducted using the software package R (R development core team 2010) with functions from the library vegan, a package for plant community ecology (Oksanen et al. 2010). RESULTS AND DISCUSSION One hundred and twenty-six vascular plant species from seventy-five genera and twenty-eight families were identified from the alpine zones of Mt. Washburn. The species list includes one fam- ily and one genus from Lycophyta, two families, four genera, and four species of Gymnosperms, and twenty-five families, seventy genera, and one hundred twenty-two species of Angiosperms. Important families included Asteraceae (24 spe- cies), Brassicaceae (14 species), Poaceae (14 species), Cyperaceae (6 species), Polygonaceae (6 species), and Scrophulariaceae (6 species). Washburn Communities Aho and Weaver (2010) used cluster and pruning analysis to objectively identify six nodal communi- ty types on Mt. Washburn. These included two turf 2012] communities (dense turf and rocky turf), along with ridgetop, snowbank, talus, and ledge types. This paper adds two other general Mt. Washburn associations: treelines and disturbed environments. Descriptions of these communities, which follow, include comparisons to similar communities re- ported for our region (Northern-Central Rockies), the larger Rocky Mountains region, and coastal Cordilleras. A primary objective of this survey was to compare the flora of Mt. Washburn to those of other North American andesitic-alpine locations. Dense turf. North-facing slopes on Washburn were characterized by dense dry meadows dom- inated by Carex elynoides Holm, secondary graminoids including Carex obtusata Lilj., Luzula spicata (L.) DC., and Poa glauca Vahl var. rupicola (Nash ex Rydb.) Boivin, and perennial forbs including Minuartia obtusiloba (Rydb.) House, Cerastium arvense L., Polemonium visco- sum Nutt., Potentilla diversifolia Lehm.var. diver- sifolia, and Sedum lanceolatum Torr. Dry Carex elynoides turf is ubiquitous to the Rocky Mountain alpine from Montana (Bamberg and Major 1968; Cooper et al. 1997; Damm 2001; Aho 2006) through Idaho (Caicco 1983; Urbanczyk and Henderson 1994; Richardson and Henderson 1999), Utah (Lewis 1970), Wyoming (Billings and Bliss 1959), Colorado (Komarkova and Weber 1978; Komarkova 1979; Willard 1979; Hartman and Rottman 1988), and New Mexico (Baker 1983). On andesitic substrates C. elynoides turf occurs locally in the northern (Aho 2006) and southern Absarokas (Thilenius and Smith 1985), and at most other documented andesitic/alpine ranges in the Rockies including Buffalo Peaks and San Juans of southern Colorado (Rottman and Hartman 1985; Seagrist and Taylor 1998), and the Sangre de Cristo Mountains in New Mexico (Baker 1983). Notably, Trifolium dasyphyllum Torr. & A. Gray is co-dominant with Carex elynoides on andesitic substrates in the southern Absarokas (Thilenius and Smith 1985) and on Latir Mesa in the Sangre de Cristo Mountains (Baker 1983). These species also co-occur on both granitic and limestone substrates in the nearby Beartooths (Aho 2006). The genus Trifolium L., however, does not occur in the alpine of Mt. Washburn, or on the northernmost peaks of the nearby andesitic Northern Absarokas (Aho 2006). Rocky turf. Rocky turf communities occupied heterogeneous patches on steep, south facing slopes. These were often dominated by Packera cana (Hook.) W. A. Weber & A. Léve, and Astragalus kentrophyta A. Gray var. tegetarius (S. Watson) Dorn, with associates Minuartia obtusi- loba, Cerastium arvense, Erigeron compositus Pursh var. discoideus A. Gray, Lomatium cous AHO AND BALA: ALPINE FLORA OF MOUNT WASHBURN 5 (S. Watson) J. M. Coult & Rose, Phlox pulvinata (Wherry) Cronquist, and Sedum lanceolatum. This association is similar to windswept dry non- prostrate communities in the Tendoy and Tobacco Root Mountains in southwest Montana dominated by Lomatium cous, Phlox pulvinata, Sedum lanceolatum and Smelowskia calycina (Steph. ex Willd.) Meyer var. americana (Regel & Herder) W.H. Drury & Rollins (Cooper et al. 1997). On andesitic substrates an Erigeron compositus- Astragus kentrophyta cushion plant community occurs locally in northern Absarokas (Aho 2006). A similar Packera cana-A. kentrophyta-E. compo- situs association occurs in andesitic rocky envi- ronments in the Sweetwater Mountains in the Sierra Nevada (Hunter and Johnson 1983). Ridgetop environments. Ridgetops were often dominated by Erigeron rydbergii Cronquist, Oxy- tropis lagopus Nutt., and cushion plants species, including Minuartia obtusiloba, Astragalus ken- trophyta, Phlox pulvinata, Draba densifolia Nutt., Draba incerta Payson, Eriogonum ovalifolium Nutt., Erigeron compositus, and Selaginella densa Rydb. Erigeron rydbergii 1s endemic to the Greater Yellowstone Ecosystem, and is limited in distri- bution to southern Montana and northwestern Wyoming and parts of Idaho (Pemble 1965; Hitchcock and Cronquist 1973; Lackschewitz 1994). Selaginella densa and Erigeron compositus frequently co-occur in cushion plant communities in the Pioneer, Tobacco Root, Madison, Beaver- head and Tendoy Ranges in southwestern Mon- tana (Cooper et al. 1997), although these associations often include and Dryas octopetala L.var. hookeriana (Juz.) Breitung and Geum rossii (R. Br.) Ser. Both G. rossii and D. octopetala are absent from Mt. Washburn. Astragalus kentro- phyta, Draba densifolia, Erigeron compositus, and Phlox pulvinata occur frequently on rocky an- desitic substrates in the Sweetwater Mountains in the Sierra Nevada (Hunter and Johnson 1983). Talus and scree. Elymus scribneri (Vasey) M. E. Jones frequently dominated heterogeneous patch- es in talus with Erigeron compositus and Ceras- tium arvense. Other infrequent associates included Chaenactis alpina (A. Gray) M. E. Jones, Carex haydeniana Olney, and Polemonium viscosum. Species composition on scree is similar to that at other north-central Rocky Mountains loca- tions. In particular, rocky areas in southwestern Montana are often dominated by Elymus scrib- neri, Festuca brachyphylla Schult. & Schult. var. coloradensis (Fred.) Dorn, Trisetum spicatum (L.) K. Richt., Achillea millefolium L. var. lanulosa (Nutt.) Piper, and Lomatium cous (Cooper et al. 1997). Rocky grassland communities of the Copper Basin in Idaho are dominated by E/ymus scribneri, Poa glauca var. rupicola, and Erigeron compositus (Caicco 1983). While E. scribneri is G MADRONO widespread from Utah (Hayward 1952; Lewis 1970), to Colorado (Hartman and Rottman 1988) to Montana (Pemble 1965) and New Mexico (Hitchcock and Cronquist 1973), it has only been noted as a major alpine community component in the northern and north-central Rocky Mountains (e.g., Caicco 1983; Cooper et al. 1997). With regard to andesitic substrates Elymus scribneri and Erigeron compositus frequently co- occur in the northern (Aho 2006) and southeast- ern Absarokas (Thilenius and Smith 1985). Elymus scribneri occurs mostly in dry meadows in the andesitic Buffalo Peaks of southern Colorado (Seagrist and Taylor 1998). Snowbank environments. Snowbank areas on Mount Washburn were dominated by Carex pay- sonis Clokey and Artemisia scopulorum A. Gray. Secondary species included the graminoids Carex phaeocephala Piper, Festuca brachyphylla var. coloradensis and Luzula spicata, and the forbs Achillea millefolium var. lanulosa, Minuartia ob- tusiloba, Cerastium arvense, Erigeron simplex Greene, Polygonum bistortoides Pursh, Sibbaldia procumbens L., and Stellaria monantha Hulten. Locally this association appears to be similar to several Carex paysonis snowbank communities on sedimentary ranges of southwestern Montana (Cooper et al. 1997). Carex paysonis communities also occur in the granitic Beartooths (Aho 2006), and on neo-glacial deposits in the Tetons (Spence and Shaw 1981). Carex paysonis associations appear frequently on andesitic substrates in the Rocky Mountain and coastal cordilleras. Carex paysonis- Artemisia scopulorum l\ate-melt communities occur locally on peaks in the andesitic northern Absarokas (Aho 2006). Similar associations also occur on moist and wet andesitic meadows in the distant Buffalo Peaks (Seagrist and Taylor 1998), and in the San Juan and Sangre de Cristo Mountains in the Southern Rockies (Baker 1983; Rottman and Hartman 1985). Wet meadows in the San Juan Mts. include A. scopulorum, Erigeron simplex, Sedum integrifolium (Raf.) A. Nelson, and Sib- baldia procumbens (Rottman and Hartman 1985), while similar sites in Sangre de Cristo Mountains include A. scopulorum, Lloydia_ serotina (L.) Rchb., and Salix arctica Pall. var. petraea (Andersson) Bebb (Baker 1983). Carex paysonis late melt communities also occur on volcanic Mount St. Helens in Washington (del Moral and Jones 2002) and on Mount Hood in Oregon (Titus and Tsuguzaki 1999), Ledges under cliff formations. The upright forb Arnica rydbergii Greene frequently dominated runnels along cliff bases, and unstable, steep, rocky volcanic slopes. Arnica rydbergii often occurs as a patchy monoculture, although infre- quent associates include Elymus scribneri, and Cirsium eatonii (A. Gray) Robins. [Vol. 59 The northern Rockies support other similar Arnica associations. A community dominated by Arnica diversifolia Greene, Epilobium anagallidi- folium Lam., Poa alpina L., and Poa cusickii Vasey var. pallida (Soreng) Dorn pioneers wet, rocky, recently deglaciated substrates in Glacier National Park (Damm 2001). Arnica longifolia Eaton, Poa reflexa Vasey & Scribn. ex Vasey, and Ranunculus eschscholtzii Schlecht. dominate sub- alpine ledges and draws in Grand Teton National Park (Gregory 1983). Monoculture stands of Arnica sp. occur in rocky cirques, and under lime- stone outcrops of the Bridger Range of southern Montana (S. Forcella unpublished data). With regard to andesitic substrates, Arnica rydbergii-Epilobium clavatum Trel. communities often dominate low alpine cliff runnels in the northern Absarokas (Aho 2006), while A. rydber- gii occurs at alpine/subalpine ecotonal environ- ments in the volcanic northern Cascades (Douglas and Bliss 1977). Disturbed environments. Despite the general resistance of alpine areas to weed invasion (Billings and Mooney 1968), six exotic species were collected within the Washburn alpine. These included a rhizomatous grass (Bromus inermis Leyss.), a taprooted perennial forb (Taraxacum officinale Weber), and annual/biennial forbs (Polygonum aviculare L., Chenopodium rubrum L., Descurainia sophia [L.] Webb ex Prantl, and Lepidium sp. L.). It should be emphasized that these species persisted not only at subalpine- alpine ecotonal elevations, but also in areas far above treeline. The number of non-natives is notable given exotic species reports in other alpine studies. For instance, only one species (Artemisia biennis Willd.) out of 173 was reported to be exotic in the alpine regions in the Mosquito Range in central Colorado (Seagrist and Taylor 1998), while several alpine species lists report no exotics whatsoever (e.g., Spence and Shaw 1981; Baker 1983; Hunter and Johnson 1983; Hartman and Rottman 1988). The large number of exotics on Washburn is surely due to invasion vectors pro- vided by frequent human visitors and associated disturbance at trails and roadsites (cf. Weaver et al. 2001). With the exception of 7. officinale, which also inhabited wet turf sites, exotic species were generally limited to areas within and along- side roads, and other areas of heavy anthropo- genic disturbance. Treeline environments. Because treelines on Washburn are the result of historical patterns of forest fires as well as altitudinal and topographic gradients (Barrett 1994; Peet 2000), it was often difficult to distinguish alpine and high-subalpine ecotones. Species distributions were also inade- quate in distinguishing the zones. For instance, Eriogonum flavum Nutt., Delphinium bicolor 2012] — p09) SS --- N. Absarokas fo) Beartooths 2h. — Washburn ” ” 2 2) 6 oe ro) = ro) e re) x © 5 8 oO i oO ie) 5 10 15 20 25 30 35 Plots FIG. 2. AHO AND BALA: ALPINE FLORA OF MOUNT WASHBURN i (b) --- N. Absarokas Beartooths — Washburn Log (abundance) 0 20 40 60 80 100 120 140 Species rank in abundance Biogeographic analyses for the Washburn Range, nine mountains in the N. Absaroka-Volcanics, and six mountains on the Beartooth Plateau. (a) Species area curves from a first order jackknife procedure. Gray lines are 95% confidence intervals. (b) Rank abundance dominance (RAD) curves. Abundance responses were fit to a log- Normal distribution (Preston 1948). Nutt., and Geum triflorum Pursh dominated high subalpine slopes, while being absent from defin- itive alpine areas (and so are not included in this list). Conversely, other species, such as Besseya wyomingensis (A. Nelson) Rydb., Poa pattersonii Vasey, Erigeron compositus, Androsace septentrio- nalis L.var. subulifera A.Gray, Antennaria micro- phylla Rydb., Agoseris glauca (Pursh) Raf. var. dasycephala (TYorr.& A. Gray) Jeps., and Achillea millefolium var. lanulosa had broad altitudinal distributions and frequently occurred at both the highest altitudes and at areas far below treeline (and are included here). Species which were representative of the treeline ecotone included Arnica latifolia Bong., Linum lewisii Pursh, Phleum alpinum L., Penstemon attenuatus Doug- las ex Lindl., and Vaccinium scoparium Leiberg ex Coville. Ecotonal tree species (1.e., Pinus albicaulis Engelm., Picea engelmannii Parry ex Engelm. and Abies lasiocarpa (Hook) Nutt. are included in the species list to indicate the species which demark the treeline. Mt. Washburn in a Biogeographic Context Results from biogeographic analyses indicate that, for the region, Mt. Washburn has both lower levels of richness (Fig. 2a), and lower levels of immigration (Fig. 2b). In particular, Washburn was predicted to have 78 species per 36 plots, while the Beartooths and northern Absarokas were predicted to have 97 and 123 species per 36 plots. Lower immigration rates can be deduced by the sharp decline and the end of the Washburn rank abundance curve in Fig. 2b, indicating few rare species (Hubbell 2001). Both of these results fit with predictions of the theory of island bio- geography for small, isolated environments (MacArthur and Wilson 1963, 1967). That Mt. Washburn may be affected by its island charac- teristics gives rise to management concerns, particularly given recent climate models for Yellowstone National Park. These models gener- ally predict an increase in treeline elevation, further decreasing alpine island area, and increas- ing fragmentation (Romme and Turner 1990). As a result of its depauperate flora, a large number of species are missing from the Mt. Washburn alpine that are common to surrounding alpine regions. These include: Agrostis variabilis Rydb., Bupleurum americanum J. M. Coult. & Rose, Carex scirpoidea Michx., Eritrichium nanum (Vill.) Schrad. ex Gaudin var. e/ongatum (Rydb.) Cronquist, Deschampsia cespitosa (L.) P. Beauv., Dryas octopetala, Geum rossii, Pedicularis groen- landica Retz., Silene acaulis (L.) Jacq.var. sub- acaulescens (F. N. Williams.) Fernald & H. St. John, and the genus Trifolium (e.g., T. dasyphyl- lum, T. haydenii Porter, and T. parryi A. Gray). Hypothetically these absences may be due to se- veral factors including Mt. Washburn’s small size and isolation (discussed above), and its andesitic- volcanic substrate. Two of the missing species are documented calciophiles: Dryas octopetala (Bam- berg and Major 1968; Komarkova 1979; Willard 1979), and E. nanum var. elongatum (Bamberg and Major 1968). D. octopetala, B. americanum, and Eritrichium nanum var. elongatum are also 8 MADRONO absent from andesitic alpine areas in the southern Rockies (Baker 1983; Rottman and Hartman 1985) and the coastal Cordillera (Hunter and Johnson 1983). All three species however occur on southern peaks of the andesitic northern Absar- okas (Hartman et al. unpublilshed). Alpine species in the genus Trifolium (e.g., T. dasyphyl- lum, T. haydenii, and T. parryi) are also absent from the alpine of the northernmost peaks of the nearby andesitic northern Absarokas (Aho 2006). Trifolium, however, is present on andesitic substrates of southern peaks in the northern Absarokas (Rosenthal 1999; Hartman et al. upublished), and in the southern Absarokas (Thilenius and Smith 1985; Rosenthal 1999). Little evidence exists to link other missing species to substrate effects. ANNOTATED SPECIES LIST FOR THE MOUNT WASHBURN ALPINE The annotated list which follows includes species names along with qualitative information about constancy, local abundance, moisture regime, habitat preference, and native/exotic status. The constancy of each species (1.e., rare = rare, unco = uncommon, comm = common, wide = widespread), is reported first in the annotated list. Constancy records the tendency of a species to occur in all possible examples of its preferred habitat. Local abundance (i.e., scarce = scarce, abund = abundant, dom = dominant) reflects species dominance within its preferred habitat. Soil water preference (1.e., dry, moist, wet) was quantitatively determined, using soil sensors, for 59 species growing within >5% of plots examined by Aho and Weaver (2010); also see Aho (2006, Chapter 2). Water preference was subjectively evaluated for species found outside of plots. General habitat preference is denoted as: dtf = dense turf, rtf = rocky turf, rt = ridgetop, ta = talus, sm = late melt, d = disturbed, tr = treeline, / = ledges, a// = all habitats. Constancy, abundance, and water preferences are not inferred for species unless they were modeled by Aho (2006). It should be acknowledged that while the list of species in this paper is based on a large number of current and historical collections, additional rare species may still be found (J. Whipple, Yellow- stone National Park, personal communication). Sull other unlisted species may exist intermittently in the Washburn alpine as a product of random ecological drift (Hubbell 2001). LYCOPHYTA Selaginellaceae Selaginella densa Rydb. [Aho 303 YELLO]; wide, abund, dry, dtf, rtf, rt, native [Vol. 59 ANTHOPHYTA-MONOCOTYLEDONEAE Cyperaceae Carex albonigra Mack. [Gentholts, D. YELLO 4702|; wet, ta , native Carex elynoides Holm [Aho 152, 381 YELLO}; wide, dom, dry, dtf; rt, native Carex haydeniana Olney [Gentholts, D. YELLO 4653]; unco, abund, wet, sm, native Carex obtusata Lilj. [Aho 146, 335 YELLO}; comm, dom, dry, dtf, native Carex paysonis Clokey; [Aho 145, 380 YELLO]; comm, dom, wet, sm, native Carex phaeocephala Piper [Aho 144 YELLO}]; unco, abund, moi to wet, sm, d, native Juncaceae Juncus drummondii E. Mey. [Aho 544 YELLO]; unco, abund, wet, sm, native Luzula spicata (L.) DC. [Aho 564 YELLO]; wide, abund, wet to dry, a/, native / Liliaceae Allium cernuum Roth [Aho 108 YELLO]; unco, abund, dry to moist, d, tr, native Poaceae Bromus inermis Leyss. var. inermis [Aho 94 YELLO]; unco, dom, dry to moist, d, exotic Elymus scribneri (Vasey) M. E. Jones [Aho 557 IDS]; comm, abund, dry to wet, fa, rt, native Elymus trachycaulus (Link) Gould ex Shinners var. andinus (Scribn. & J. G. Sm.) Dorn [Aho SS YELLO]; unco, abund, dry to wet, d, exotic Festuca brachyphylla Schult. & Schult. var. coloradensis (Fred.) Dorn [Aho 576 IDS]; wide, abund, dry to wet, a//, native Phleum alpinum L. [Aho 567 IDS]; unco, abund, wet to moist, tr, d, native Poa alpina L. [Aho 541 IDS]; comm, abund, dry to wet, dtf, sm, native Poa cusickii Vasey var. pallida (Soreng) Dorn [Aho 360, 85 YELLO]; wide, abund, dry to wet, all, native Poa cusickii Vasey var. epilis (Scribn.) C. L. Hitche. [Aho 80 YELLO]; wide, abund, dry to wet, all, native Poa glauca Vahl var. rupicola (Nash ex Rydb.) B. Boivin [Aho 75, 382 YELLO]; comm, abund, dry to wet, dif , rtf, native Poa interior Rydb. [Aho 379 YELLO]; unco, scarce, wet to moist, fa, sm, native Poa pattersonii Vasey [Aho 77 YELLO]; wide, abund, dry to wet, dtf, rtf , native Poa reflexa Vasey & Scribn. ex Vasey [Aho 549 IDS]; unco, scarce, wet, sm, native 2012] Poa secunda J. Pres var. incurva (Scribn. & T. A. Williams. ex Scribn.) Beetle [Aho 72 73 YELLO]; unco, abund, dry to moist, ta, native Trisetum spicatum (L.) K. Richt. [Caprio, T. YELLO 4588]; wide, abund, dry to wet, a//, native ANTHOPHYTA-DICOTYLEDONEAE Apiaceae Lomatium cous (S. Watson) J. M.Coult. & Rose [Aho 559 IDS]; wide, abund, dry to wet, all, native Asteraceae Achillea millefolium L. var. lanulosa (Nutt.) Piper [Aho 271 YELLO]; wide, abund, wet to dry, all, native Agoseris glauca (Pursh) Raf. var. dasycephala (Torr. & A. Gray) Jeps. [Aho268 YELLO]; wide, abund, dry to wet, a//, native Antennaria media Greene; [Aho 482 YELLO]; unco, abund, wet, sm, native Antennaria microphylla Rydb. [Aho 577 IDS]; wide, abund, dry to wet, a//, native Antennaria umbrinella Rydb. [Aho 565 IDS]; wide, abund, dry to wet, a//, native Arnica latifolia Bong. [Aho 572 IDS]; unco, abund, wet, sm, tr, native Arnica longifolia D. C. Eaton [Condon, D. YELLO 2688]; Frequent, tr, native Arnica rydbergii Greene [Aho 558 IDS]; comm, dom, wet, fa, /, d, tr, native Artemisia scopulorum A. Gray [Aho 252 YELLO}]; unco, abund, wet, sm, native Chaenactis alpina (A. Gray) M. E. Jones var. alpina [Aho 244 YELLO]; unco, abund, dry to moist, fa, d, native Cirsium eatonii (A. Gray) B. L. Rob. [Aho 243 YELLO]; unco, abund, dry to moist, ta, /, d, native Ericameria suffruticosa (Nutt.) G. L. Nesom [Aho 228 YELLO}]; unco, abund, dry to moist, ta, rt, d, native Erigeron compositus Pursh var. discoideus A. Gray; comm. [Aho 556 IDS]; abund, dry, rz, ta, native Erigeron rydbergii Cronquist [Aho 234, 378 YELLO]; Frequent, dry, dtf, rt, native Erigeron simplex Greene [Aho 233 YELLO}]; Frequent, dry to wet, sm, dtf , native Erigeron ursinus D. C. Eaton [Currie, M C. YELLO 4265]; native Oreostemma alpigenum (Torr. & A. Gray.) Greene var. haydenii (Porter) Nesom. [Aho 249 YELLO]; unco, abund, dry to moist, sm, dtf, native (was Aster alpigenus) AHO AND BALA: ALPINE FLORA OF MOUNT WASHBURN 2 Packera cana (Hook.) W. A. Weber & A Léve [Aho 224 YELLO]; comm, abund, dry to moist, fa, rf, native Packera subnuda (DC.) Trock & T. M. Barkley [Caprio, T. YELLO 4402]; native Senecio fremontii Torr. & A. Gray [Aho 220 YELLO]; comm, abund, dry to moist, ta, rt, native Senecio integerrimus Nutt. var. integerrimus [Aho 218 YELLOJ]; unco, abund, moist to wet, sm, tr, native Solidago multiradiata Aiton var. scopulorum A. Gray [Aho 212 YELLO]; comm, abund, dry to moist, fa, d, native Symphyotrichum foliaceum (Lindl. ex DC.) G. L. Nesom var. apricum (A. Gray) G. L. Nesom [Aho 248 YELLO]; comm, abund, wet, sm, native (was Aster foliaceus var.apricus) Taraxacum ceratophorum (Ledeb.) DC. [Conrad, H. S. YELLO 3193]; comm, abund, dry to moist, a//, native Taraxacum eriophorum Rydb. [Aho 210 YELLO]; rare, scarce, moist, fa, native Taraxacum officinale Weber [Aho 208 YELLO]; comm, abund, moist to wet, d, exotic Townsendia parryi Eaton [Aho 207 YELLO}]; tr, native Boraginaceae 205 dtf, Mertensia alpina (TYorr.) G. Don [Aho YELLO]; comm., abund, dry to moist, rtf, sm, native Myosotis alpestris F. W. Schmidt [Aho 204 YELLO]; comm., abund, dry to moist, dtf, rtf, sm, native Brassicaceae Boechera angustifolia (Nutt.) Dorn [Smith, F. H. YELLO /203]; unco, abund, dry to moist, fa, d, native Boechera exilis (A. Nelson) Dorn [Aho 199, 190 YELLO]; unco, abund, dry to moist, fa, d, native Boechera lemmonii S. Watson [Caprio,_ T. YELLO 4363]; unco, abund, dry to moist, fa, d, native Boechera lyallii (S. Watson) Dorn [Aho 547 IDS]; unco, abund, dry to moist, dt/, native Boechera microphylla Nutt. [Aho 568 IDS]; unco, abund, dry to moist, ta, d, native Descurainia sophia (L.) Webb ex Prantl [Aho 562 IDS]; unco, abund, dry to moist, d, exotic Draba cana Rydb. [Aho 189 YELLO]; rare, abund, dry to moist, ta, tr, native Draba crassifolia Graham [Aho 324 YELLO}]: comm, scarce, moist to wet, sm, native Draba densifolia Nutt. [Smith, Fo. H. YELLO 1266]; unco, abund, dry, rf, native 10 MADRONO Draba incerta Payson [Aho 326 YELLO]; comm, abund, dry to moist, fa, rt, native Draba paysonii J. F. Macbr. var. treleasii (O. E. Schulz) C. L. Hitche. [Aho 186 YELLO}]; rare, scarce, dry to moist, fa, rt, native Lepidium sp. L. [Aho 178 YELLO]; unco, scarce, dry to moist, d, t, exotic Noccaea parviflora (A. Nelson) Holub [Aho 176 YELLO]; unco, abund, dry to moist, tr, dtf, native Smelowskia calycina (Stephan ex Willd.) C. A. Meyer var. americana (Regel & Herder) W. H. Drury & Rollins [Woolf, A. YELLO 1/317); comm, abund, dry to moist, a//, native Caryophyllaceae Cerastium arvense L. [Aho 546 IDS]; comm, dom, moist to wet, sm, dtf, rtf, ta, native Eremogone congesta (Nutt.) Ikonn. var. lithophila (Rydb.) Dorn [Aho 174 YELLO]; comm, abund, moist to wet, sm, dtf, rtf, ta, native Minuartia obtusiloba (Rydb.) House [Aho 363 YELLOJ]; comm., abund, moist to wet, sm, dtf, ta, native Minuartia rubella (Wahlenb.) Hiern [Aho 170 YELLO]; widespread, abund, dry to wet, sm, dtf, ta, native Silene kingii (S. Watson) Bocquet [Aho 1/65 YELLO]; unco, dom, moist to wet, sm, dtf, native Stellaria monantha Hultén [Aho 161 YELLO}]; unco, abund, moist to wet, sm, dtf, native Stellaria umbellata Turcez. ex Kar. & Kir. [Conrad, H. S. YELLO 1/026]; unco, scarce, wet, sm, native Chenopodiaceae Chenopodium rubrum L. [Aho 551 YELLO]; unco, abund, d, exotic Crassulaceae Sedum lanceolatum Torr. [Aho 159 YELLO]; wide, dom, dry to wet, a//, native Ericaceae Vaccinium scoparium Leiberg ex Coville [Aho 560 YELLO]; unco, abund, wet, sm, tr, native Fabaceae Astragalus alpinus L. [Conrad, H. S. YELLO 1596]; comm, abund, dry to wet, sm, dtf, native Astragalus kentrophyta A. Gray var. tegetarius (S. Watson) Dorn [Aho 578 IDS]; comm, abund, dry to moist, rt, ta, native Astragalus miser Douglas [Conrad, H. S. YELLO 1587|; unco, dom, dry to moist, tr, dtf, native [Vol. 59 Lupinus argenteus Pursh [Aho 129 YELLO]; wide, dom, dry to moist, dtf, rt, sm, native Oxytropis borealis DC. var. viscida (Nutt.) S. L. Welsh [Aho 127 YELLO]; unco, abund, dry, rt, native Oxytropis lagopus Nutt. [Aho 126 YELLO]; comm, abund, dry, dif, rt, native Oxytropis parryi A. Gray [Aho 125 YELLO}; rare, scarce, dry to moist, dtf, native Grossulariaceae Ribes montigenum McClatchie [Aho 118 YELLOJ; unco, abund, moist, tr, sm, native Hydrophyllaceae Phacelia hastata Douglas ex Lehm. [Aho 579 IDS]; unco, abund, dry to moist, ta, native Phacelia sericea (Graham ex Hook.) A. Gray [Aho 116 YELLO]; unco, abund, dry to moist, ta, native Linaceae Linum lewisii Pursh [Aho 56 I IDS]; unco, abund, dry to moist, ta, dtf, tr, native Onagraceae Epilobium clavatum Trel. [Aho540 IDS]; unco, abund, wet, ta, sm, native Epilobium halleanum Hausskn. [Aho 101 YELLO]; rare, scarce, wet, sm, native Parnassiacea Parnassia fimbriata Kk. D. Konig [Condon, D. YELLO 1/385]; wet, tr, native Polemoniaceae Phlox multiflora A. Nelson [Aho 580 IDS]; wide, dom, dry to moist, tr, dtf, rtf, native Phlox pulvinata (Wherry) Cronquist [Aho 168, 398, 399 YELLO]; comm, abund, dry, r¢, dtf, rtf, native Polemonium pulcherrimum Hook. [Aho 555 IDS]; unco, abund, dry to moist, rocky dtf, ta, native Polemonium viscosum Nutt. [Woolf, A. YELLO 2129]; comm, dom, dry to moist, dtf, ta, native Polygonaceae Eriogonum ovalifolium Nutt. [Aho 65 YELLO]; unco, abund, dry, ré, rtf, dtf, native Oxyria digyna (L.) Hill [Aho 552 IDS]; infr, abund, moist, ta, native Polygonum aviculare L. [Aho 62 YELLO]; unco, abund, d, exotic 2012] Polygonum bistortoides Pursh [Aho 12 YELLO]; comm., abund, dry to moist, dtf, native Polygonum douglasii Greene var. microspermum (Engelm.) Dorn [Aho 539 IDS]; unco, scarce, dry to moist, fa, d, native Rumex paucifolius Nutt. [Aho 56 YELLO}]; rz, ta, native Portulacaceae Claytonia lanceolata Pursh [Aho 553 IDS]; comm, abund (early spring), dry to moist, dtf, native Lewisia pygmaea (A. Gray) B. L. Rob. [Aho 368 YELLO]; unco., abund, wet to moist, dif, sm, native Cistanthe umbellata (Yorr.) Hershk. var. caudici- fera (A. Gray) Kartesz & Gandhi [Aho 545 IDS]; unco, abund, moist, ta, d, native Primulaceae Androsace_ septentrionalis L. var. subulifera A. Gray [Aho 542 IDS]; wide, scarce, dry to moist, all, native Dodecatheon conjugens Greene [Woolf, A. YELLO 2017); dtf, native Dodecatheon pulchellum (Raf.) Merr. [Aho 51] YELLO]; unco, abund, moist to wet, dtf, sm, native Ranunculaceae Delphinium bicolor Nutt. [Aho 330, 369 YELLO]; dry to moist dtf, tr, native Ranunculus eschscholtzii Schlecht. [Aho 43 YELLO]; comm, dom (early spring), moist to wet, fa, sm, native Rosaceae Potentilla diversifolia Lehm. var. diversifolia [Aho 35, 36 YELLO]; comm, dom, dry to moist, d¢f; rtf, native Potentilla ovina J.M. Macoun [Aho 32 YELLO]; comm, abund, dry, dtf, rtf, rt, native Sibbaldia procumbens L. [Aho 74 YELLO]; unco, abund, sm, dtf, native Salicaceae Salix arctica Pall. var. petraea (Andersson) Bebb [Aho 543 IDS]; unco, abund, wet, sm, native Saxifragaceae Saxifraga cespitosa L. [Conrad, H. S. YELLO 1565]; unco, scarce, wet, sm, native Saxifraga rhomboidea Greene [Aho 400, 401 YELLO]; comm, abund, dry to moist, dtf; native AHO AND BALA: ALPINE FLORA OF MOUNT WASHBURN o Scrophulariaceae Besseya wyomingensis (A. Nelson) Rydb. [Aho 582 IDS]; comm, abund, dry to moist, df, native Mimutus lewisii Pursh [Condon, D. YELLO 2490); infrequent, wet, tr, native Pedicularis cystopteridifolia Rydb. YELLO)]; infrequent, wet, s77, native Penstemon attenuatus Douglas ex Lindl. [Aho 6 YELLO]; infrequent, dry to moist, dtf, tr, native Penstemon procerus Douglas ex Graham [Aho 583 IDS]; infrequent, dry to moist, dtf, native Veronica wormskjoldii Roem. & Schult. [Aho 538 YELLO}]; infrequent, wet, s7, native [Aho 10 CONIFEROPHYTA Pinaceae Abies lasiocarpa (Hook.) Nutt. [Condon H. S. YELLO 49]; infrequent, dry to moist, tr, native Picea engelmannii Parry ex Engelm. [Aho 537 YELLO}]; infrequent, dry to moist, tr, native Pinus albicaulis Engelm. [Condon H. S. YELLO 56]; infrequent, dry to moist, tr, native ACKNOWLEDGMENTS This research was made possible in part with a grant from the National Park Service (YNP-NPS #YELL- 05116) and by assistantships from Montana State University. Thanks to M. Hektner (YNP-NPS), H. Anderson, J. Whipple (YELLO), T. Weaver, T. Seipel (MSU), and C. Seibert and M. Lavin (MONT). LITERATURE CITED AHO, K. 2006. Alpine ecology and subalpine cliff ecology in the Northern Rocky Mountains. 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Exotic plants in early and late seral vegetation of fifteen northern Rocky Mountain environments (HTs). Western North American Naturalist 61:417-427. WILLARD, B. E. 1979. Plant sociology of alpine tundra, Trail Ridge, Rocky Mountain National Park, Colorado. Quarterly of the Colorado School of Mines 74:4. MADRONO, Vol. 59, No. 1, pp. 14-24, 2012 IMPACT OF RECURRENT FIRE ON ANNUAL PLANTS: A CASE STUDY FROM THE WESTERN EDGE OF THE COLORADO DESERT ROBERT J. STEERS! AND EDITH B. ALLEN Department of Botany and Plant Sciences and Center for Conservation Biology, University of California, Riverside, CA 92521 robert_steers@nps.gov ABSTRACT Limited information exists regarding the impact of fire on annual plant composition in creosote bush scrub vegetation. The impact of recurrent fires on annual plants is even less understood. To investigate this matter, annual vegetation was sampled in a stand of creosote bush scrub in western Coachella Valley, California that had recently experienced two wildfires. The wildfires fragmented the once contiguous shrubland into three sections: unburned, once-burned, and twice-burned stands, all of which were separated by fuel breaks that contained each fire. For all three stands, annual plant cover and species richness were determined in the field, soil seed bank samples were collected and assayed in a glasshouse, and soil chemistry and physical properties were measured. We found that invasive annual grass cover was highest in the twice-burned stand and native annual plant cover was greatest in the unburned stand. Native annual species richness significantly decreased each time a stand burned resulting in low native annual plant diversity. Seed bank assays revealed that invasive annual grass germinants were orders of magnitude greater in the twice-burned stand compared with the other two stands. Lastly, soil total N, C, and soil pH were elevated in both burned stands. Overall, we found that recurrent fire can result in strong impacts to annual vegetation; however, the twice- burned stand was sampled only three years after burning while the once-burned stand was sampled 20 years after burning. Thus, longer-term fire effect studies plus replication with additional study sites are still needed to improve our understanding of how recurrent fire impacts annual plants. Key Words: Diversity, feedback, grass/fire cycle, invasive plant, richness, seed bank. Invasive grasses can alter the fire regime by increasing the frequency, intensity, extent, and seasonality of fire (Brooks et al. 2004). In creosote bush scrub vegetation of southern California, the invasive grasses are annuals that differ funda- mentally from the native annual forbs that they displace. For example, unlike most native annual plants, invasive annual grasses senesce earlier and have persistent standing biomass throughout the dry season (Brooks 1999). One problematic result of grass invasion for creosote bush scrub is longer-lasting fine fuel that connects widely spaced shrubs (Brooks et al. 2004). In addition, invasive grasses can form higher density assem- blages than native vegetation (Steers and Allen 2010), thus increasing the fuel packing ratio and consequently, fire intensity. The primary foci of previous studies examining fire in desert shrublands of the Mojave and So- noran Deserts have been on the impacts to pe- rennial species. These studies have documented reductions in cacti and long-lived shrubs, such as Larrea tridentata Coville, and increases in rela- tively short-lived perennials (O’Leary and Muin- nich 1981; McLaughlin and Bowers 1982; Brown and Minnich 1986; Alford et al. 2005; Brooks and 'Present address: National Park Service, San Fran- cisco Bay Area Network, Inventory and Monitoring Program, Bldg. 1063 Ft. Cronkhite, CA 94965 Minnich 2006; Abella 2009; Abella 2010). How- ever, most of these studies did not measure an- nual vegetation. Previous studies that have focused on fire and desert annual plants documented post fire decreases in Bromus madritensis L. subsp. rubens (L.) Husn. and increases in Schismus spp. (either S. arabicus Nees, S. barbatus [L.] Thell., or both), which are both invasive annual grasses. These studies also documented little change in the abundance of the invasive forb, Erodium cicutarium (L.) Aiton, and either an increase or decrease in native annuals, depending on the species (Cave and Patten 1984; Brooks 2002; Esque et al 2010a, b; Steers and Allen 201 1a). Native annual plant species richness has also been shown to decline in shrub under- stories after fire but no response was detected in interspace habitat (Brooks 2002). Besides these studies, little information exists on the impacts of fire on desert annual plants. Brooks et al. (2004) and Brooks and Esque (2002) warn that post fire increases in invasive annual grasses may promote recurrent fire, sensu the grass/fire cycle (D’Anto- nio and Vitousek 1992). However, documentation of the impact of recurrent fire on annual veg- etation is lacking. The goal of this study was to document the im- pact of fire on the annual plant community in creosote bush scrub that partially burned in 1988 and in 2003. The fire history of this study site 2012] resulted in an unburned, once-burned, and twice- burned stand. Our objective was to measure the response of invasive annual grasses and forbs to fire, and the impact of fire on native annual plants. We were especially interested in docu- menting the response of native annual plant spe- cies richness and diversity measures to recurrent fire, as this is relatively unknown. In addition to vegetation surveys, seed bank samples collected from the three stands were also assayed and soil parameters (nutrients, texture, and pH) were mea- sured to provide additional insight. MATERIALS AND METHODS Study Area The study site was located in Whitewater Canyon (33°56'50”"N, 116°38'43”W) on the western edge of the Colorado Desert in Riverside County, California. Vegetation along the floor of White- water Canyon was composed of desert riparian and desert dry wash communities. Creosote bush scrub occurred throughout the valley bottom, upland of the riparian and wash areas. On steeper slopes of the canyon, Encelia farinosa Torr. was dominant. Coastal scrub shrubs, such as Artemi- sia californica Less. and Salvia apiana Jeps., were occasional on north and east facing hill slopes. Within unburned creosote bush scrub of the study site, perennial vegetation was about 21% cover (Steers and Allen 2011b). Larrea tridentata con- tributed about 10% cover, Ambrosia dumosa (A. Gray) W. W. Payne 6%, Krameria grayi Rose & Painter 3%, and the following species individually contributed less than 2% cover: Psorothamnus arborescens (A. Gray) Barneby, Ephedra califor- nica S. Watson, Encelia farinosa, and cacti, in- cluding Echinocereus engelmannii (Engelm.) Lem., Opuntia basilaris Engelm. & J. M. Bigelow and O. echinocarpa Engelm. & J. M. Bigelow (Steers and Allen 2011b). The slope at the study site was 5 to 6 degrees, facing east, at an elevation of about 525 m. Soils were alluvial (NRCS 2010) and about 80% sand (this study). Annual average pre- cipitation is assumed to be between 9.5 + 5.6 SD and 29.9 + 16.4 SD cm, based on records for Palm Springs, about 19 km to the southeast, and Cabazon, about 14 km to the southwest, respec- tively (WRCC 2008). Precipitation at the Palm Springs weather station was 3.1, 4.8 and 16 cm for calendar years 2006, 2007 and 2008, respectively. Summer precipitation was negligible during these three years, which is typical for the area (WRCC 2008). Fire disturbance in this part of the Colo- rado Desert is not uncommon, and severalburned stands of creosote bush scrub have been previ- ously investigated nearby (O’Leary and Minnich 1981; Brown and Minnich 1986; Steers and Allen 201 1b). STEERS AND ALLEN: FIRE AND DESERT ANNUALS 15 Fire History Determination The fire history of the study site was determined based on stereoscope validation of fire perimeters from a series of aerial photographs of the study landscape, spanning from 1949 to 2005. Aerial photos were obtained from Riverside County Flood Control and Water Conservation District, Coachella Valley Water District, and UC River- side Science Library. The years when aerial photos were taken include the following: 1949, 1957, 1974, 1980, 1984, 1985, 1986, 1987, 1989, 1990, 1995, 1996, 1998, 2000, and 2005. For fires that occurred after the 2005 aerial photographs, additional fires were recorded from personal ob- servation. Aerial photography revealed that the year of the first fire occurred sometime between 1987 and 1989. Because fires in desert vegetation are more common following winter seasons with above average rainfall (Brooks and Matchett 2006), the wettest year, which was 1988, is re- ported as the assumed burn year. Based on per- sonal observations, the second fire occurred in the summer of 2005, following a winter of above average precipitation (WRCC 2008). Based on aerial photography, the pattern of fire at the study site transformed an area with similar creosote bush shrub cover into three stands, one of which was a 1.7 ha remnant unburned stand, a 2.7 ha once-burned stand (burned in 1988), and a 3.3 ha twice-burned stand (burned in 1988 and 2005). At the time of both fires, fuel breaks (dozer- lines) were implemented to contain each fire from spreading into adjacent areas. Therefore, differ- ences in vegetation among all three investigated stands prior to the first fire, and differences be- tween the vegetation in the once- and twice-burned stands prior to the second fire, are assumed to be negligible. Soil and Vegetation Sampling In August of 2006, six vegetation sampling plots were implemented in a stratified random design within each unburned, once-burned, and _ twice- burned vegetation stand. A sampling plot consisted of one, 7.32 m radius, modified — National Weed Management Association (mod-NAWMA) plot (Stohlgren et al. 2005). Slope and aspect were mea- sured from the center of each plot using a compass and clinometer. Soil was collected to determine nu- trient levels, physical characteristics, and to assess the seed bank. For soil nutrients, four soil samples per mod-NAWMA plot were taken to 5 cm depth with a 2.5 cm diameter corer and pooled into one composite sample per plot. The four samples were collected at the center and at three edge locations (7.32 m from plot center), at 30, 150, and 270 de- grees from plot center. For soil seed bank samples, four cores per plot were also collected within a 20 cm radius of the soil nutrient sample plugs, 16 MADRONO except 5 cm diameter cores were used instead. These cores were also pooled into one composite sample per plot. One core with the same dimen- sions used for seed bank samples was taken at the center of the plot, within a 20 cm radius of where the other soil samples were collected, for bulk density, coarse fraction (>2 mm), and soil texture measures. All soil sampled was taken at a 5 cm depth and placed in one of three plastic Ziploc bags per plot, for soil nutrient, seed bank, and physical (bulk density, coarse fraction, and soil texture) measurements, and then transported back to UC Riverside. At UC Riverside, samples taken with the 2.5 cm diameter corer were split, and 50 g were used to measure pH in a 1:1 soil:water slurry using a Fisher Scientific® Model 50 pH meter. The re- maining portion of the soil nutrient samples were then sent to the University of California, Division of Agriculture and Natural Resources Analytical Laboratory at UC Davis for carbon (C), total nitrogen (N), KCl-extractable NH4t and NO; , and texture (% sand, silt and clay) analyses (http:// groups.ucanr.org/danranlab). Soil seed bank samples were assayed by grow- ing them out in a glasshouse and counting the number of germinants per species (Cox and Allen 2008). First, a composite sample was sieved through a 6 mm X 6 mm mesh to remove coarse materials, making sure not to remove any seeds, and then spread out on a 20 cm X 20 cm Sty- rofoam tray. Soil depth in each tray ranged from 1 cm to 2 cm. Then, trays were kept moist and germinants were removed when identifiable or at a stage where they could be transplanted safely to pots to await identification. Watering continued in all trays until no new seeds germinated and then trays were left to dry. Once the soil in each tray was completely dry, it was mixed before the next watering cycle. Three cycles of watering and drying took place from September 2007 to May 2008. Trays were allowed to dry from 3 to 6 weeks between watering periods. By the third cycle, negligible numbers of seeds germinated so further cycles were not implemented. Throughout the watering, trays were reorganized several times to minimize localized effects within the glasshouse. During the winter wet-season of 2006—07, in- sufficient rainfall prevented the germination of annual plants at the study site, and no vegetation measurements were taken. In the winter of 2007— 08, precipitation was about average and vegeta- tion was sampled in March 2008 during peak- flowering. In each established mod-NAWMA plot, percent cover by species was measured in three 1 m* (1 m X | m) quadrats, located 4.57 m from plot center at 30, 150, and 270 degrees. Species richness was measured within each of the three | m* quadrats per plot and also within each plot (out to a 7.32 m radius from plot center). All species names follow nomenclature in Hickman (1996). [Vol. 59 Data Analyses Annual plant composition between the three stands (unburned, once-burned, and _ twice- burned) was compared using presence/absence data for all annual species recorded in the six mod- NAWMA plots per stand. Plots were ordinated with Nonmetric Multidimensional Scaling using a Sorenson distance measure (McCune and Mefford 2006). A random starting configuration with 50 runs of real data was used in the autopilot mode with medium speed. Then, a Multiresponse Per- mutation Procedure, using a Sorenson distance measure, was performed on the same data to determine if there were any significant pairwise difference between annual species composition in the three stands at a = 0.05 (McCune and Mefford 2006). Other comparisons between the three stands were also conducted based on categorizations of species (e.g., native versus invasive), species rich- ness, and species similarity indices. Annual vegeta- tion cover was categorized into invasive grass, invasive forb, total invasive annuals (grass + forb), and native annuals (grass + forb) at the 1 m? scale (in quadrats). Native annual grasses and forbs were combined and not treated separately because the number of native annual grass species was very low (Appendix 1). Species richness of native an- nuals was calculated at both the quadrat and mod- NAMWA plot scales while species richness of herbaceous perennials and shrubs were only calculated at the mod-NAMWA plot scale. To determine the impact of fire on seed banks, ger- minants were grouped into four categories: invasive grasses, invasive forbs, native annuals, and shrubs plus cacti. No herbaceous perennials were found in the seed bank assays. Shannon Diversity (H’) was calculated at the quadrat scale based on native annual richness and cover by species (Shannon and Weaver 1963). Also, within-plot native annual plant species sim- ilarity (S) was also calculated from the three | m7? quadrats per mod-NAMWA plot. This was done using a multiple-quadrat community coefficient based on a modification of the Sorenson index (Diserud and Odegaard 2007): S = (3/2)([ab + ac + be — abc\/[a + b + c]), where a is the number of species in quadrat (plot frame) A, b is the number of species in quadrat B, etc... and ab, ac, bc and abe are the number of species shared between quadrats A and B, A and C, B and C and A, B and C, respectively. To improve the normality of data, germinant density and native annual plant richness were square-root transformed. One-way ANOVA and Fisher’s LSD test were used to compare the soil and vegetative variables among unburned, once- burned, and twice-burned conditions at « = 0.05. Vegetation parameters that were constituted of more than one sub-sample per mod-NAWMA 2012] TABLE 1. STEERS AND ALLEN: FIRE AND DESERT ANNUALS 17 AVERAGE SOIL AND PERENNIAL PLANT PARAMETERS FOUND IN UNBURNED, ONCE-BURNED (1988), AND TWICE-BURNED (1988 AND 2005) STANDS. For each parameter, F test statistics based on one-way ANOVA are shown. Differences in superscript letters indicate significant differences between stands based on post-hoc LSD tests. Parameters that did not differ significantly between paired stands do not contain letter superscripts. n = 6 and a = 0.05 for all statistical analyses. Unburned Parameters (avr. + SE) SOIL Total N (%) 0.08 + 0.01% Total C (%) 0.75 + 0.094 NH, (ppm) 10a cng De, NO; (ppm) i 19 Sand (%) Ji8= I Silt (%) Ly = 07 Clay (%) 52 205" pH 74+ 0.14 Bulk density (g/cm*) 1.29 + 0.09 Coarse fraction (g/cm*) 0.31 + 0.03 Bare ground cover (%) Polite noe Rock cover (%) 34 = 15° Litter cover (%) 3.7 Se liad HERBACEOUS PERENNIALS Richness (species/168.3 m7?) 0.8 + 0.4 SHRUBS Richness (species/168.3 m*) 6= 05° Live cover (%) 10.6 + 3 Encelia farinosa cover (%) = o> plot (e.g., percent cover and species richness per m°) were averaged together before analyses. Thus, for all soil and vegetation analyses, n = 6. Similar Statistical procedures have been utilized when comparing paired burned and unburned vegeta- tion in low replication contexts (Haidinger and Keeley 1993; Brooks and Matchett 2003; Abella et al. 2009). RESULTS Impact of Fire on Soils Some soil parameters were influenced by fire while others were not (Table 1). Extractable nitrogen (NH,* and NO;_) did not differ between paired burned and unburned areas. However, total N and C, and soil pH were greater in the twice- burned area than the unburned area. Also, percent cover of bare ground and rock were greatest in the twice-burned area (Table 1). Impact of Fire on Seed Banks A total of 6357 germinants belonging to 14 spe- cies were recorded from the seed bank study. About 97% of the seed bank germinants were Schismus barbatus and S. arabicus, which are col- lectively referred to as Schismus spp. Invasive forbs, Erodium cicutarium and Brassica tourne- fortii Gouan, made up about 0.6%, with 29 and 12 individuals counted, respectively. Only four Bro- mus madritensis ssp. rubens individuals were Once-burned Twice-burned (avr. + SE) (avr SE) F 0.10 + 0.0148 0.13 + 0.018 61:15 0.96 + 0.0748 | ee memos Oe be 6.8488 13:7 = TA [42 = al 0.1046 Leo a= We2 14.6 24 0.9811 80 + 1.1 80.5 = 0.8 2.1964 16.7 = 15 2207 1.8023 33 = 04 As = 0.7 5.6707 7.6 =O? age ae 9.1424 1.15 + 0.06 1.15 + 0.08 1.0887 OAD O13 0.49 + 0.03 1.3802 23:53 18.9 + 3.48 2.4091 57 I21= 2:5" 4.8556 5.0 22 29 16+ 0.6 1.3029 0O+0 05 = 03 1.9000 pAak! 0 as 2 =. 028 46.8182 18.6 + 9.3 18.6 + 5.1 0.5323 [79.9024 16.4 + 4.88 2.6668 counted, all of which only occurred in soils from the unburned area. Eight native annual species made up about 1.8% of the total germinants counted. The most abundant native species were Camissonia californica (Torr. & A. Gray) P. H. Raven, Crassula connata (Ruiz & Pav.) A. Berger, and Plantago ovata Forssk. Only Encelia farinosa and one unknown cactus that died prematurely made up the six germinants in the shrub plus cacti category. Analyses of the seed bank at a scale of 78.5 cm? showed that the twice-burned stand had greater invasive annual grass density than the once- burned and unburned stands (Fig. 1). No differ- ences in invasive forb and native annual germi- nants were found between stands (Fig. 1). At this small scale, mean native annual plant richness also was not different between the unburned (1.33 + (0.33 SE species), once-burned (1 + 0.52 SE), and twice-burned (1.33 + 0.33 SE) stands. Impact of Fire on Aboveground Vegetation Five exotic annuals and 38 native annual species (Appendix 1) were documented. Of the exotics encountered, the invasive forbs, Brassica tourne- fortii and Erodium cicutarium, were widespread in all three areas, as were the invasive grasses, Schismus spp. When comparing annual plant spe- cies composition among the three paired stands, the Nonmetric Multidimensional Scaling (NMS) analysis resulted in a two-dimensional solution. Lg MADRONO L] Unburned L] Once-Burned a) Invasive Grass Density _ b) Invasive Forb Density 1000 2 _ 5 @ 900 2a 4 _— ia = gs a3 ES 2007. Bo 2 cy 100 O 1 0 0 Fic. 1. [Vol. 59 M Twice-Burned c) Native Annual Density a 14 —° 12 a sa) Germinants/ 78.5 cm? Average density and SE bars for invasive grass (a), invasive forb (b), and native annual (c) germinants from seed bank assays of the unburned, once-burned, and twice-burned stands. Differences in letters above bars indicate significant differences between stands based on ANOVA and LSD tests (a = 0.05). The final stress for the best solution was low, at 12.23 out of 100. The proportion of variance represented by each axis, based on the r* between distance in the ordination space and distance in the original space was 0.681 and 0.209 for axis 1 and 2, respectively. Thus, the separation that was revealed among the three stands was primar- ily along axis 1 (Fig. 2). Species that had the strongest correlation with axis 1 were Bromus madritensis ssp. rubens (r = —0.851), Plantago ovata (r = —0.851), Chaenactis fremontii A. Gray (r = —0.841), Vulpia octoflora (Walter) Rydb. (r = —0.819), Pectocarya linearis DC. (r = —0.696), Stylocline gnaphaloides Nutt. (r = —0.658), Pholistoma membranaceum (Benth.) Constance (r = —0.653), and Phacelia distans Benth. (r = —0.652). When the three stands were compared using the Multi-Response Permutation Procedure, the unburned stand had a significantly different annual plant community than the once- burned (A = 0.296; P < 0.001) and twice-burned (A = 0.299; P < 0.001) stands. Similarly, the , Axis 2 -1.5 -0.5 0.5 1.5 Axis 1 Fic. 2. A Nonmetric Multidimensional Scaling ordi- nation of unburned, once-burned, and _ twice-burned sample units (white, grey, and black triangles, respec- tively) based on presence/absence of annual species recorded in 168.3 m*? mod-NAWMA plots. once-burned and twice-burned stands were also significantly different from each other (A = 0.134; P= 0001), No difference in total invasive annual plant cover between the unburned, once-burned and twice-burned stands was detected (Fig. 3). How- ever, invasive grass cover was greater and invasive forb cover was lower in the twice-burned compared to the once-burned stand (Fig. 4). In addition, relative cover of invasive grasses was greatest in the twice-burned stand versus the other two stands (Fig. 3). Native annual plant cover and species richness at the 1 m* scale were lower in both of the burned stands compared to the unburned stand (Figs. 3 and 4). Also at the 1 m? scale, the stand that had burned twice did not have lower native cover or richness compared to the stand that had only burned once. However, native annual plant diversity (Shannon Diversity — H') was lower in the twice-burned stand com- pared to the other two stands (Fig. 3). Also, only in the twice-burned stand was within-plot native annual plant similarity (based on shared species among the three 1 m’ quadrats per plot) greater than the unburned stand (Fig. 3). In other words, the variety of annual species found in twice- burned vegetation was lower compared to the unburned stand. At a larger scale (168.3 m* mod- NAWMA plot), native annual species richness was lower within each burned stand (Fig. 3). Also at this larger scale, shrub richness was lower for the first burn, but showed no further decrease after the second burn. Herbaceous perennial rich- ness was very low in general and did not differ among the three stands (Table 1). DISCUSSION Impact of Fire on Soils Soil pH, and total N and C were greatest in the twice-burned stand, which last experienced fire three years prior to sampling. Elevated pH is 2012] STEERS AND ALLEN: FIRE AND DESERT ANNUALS 19 [_]Unburned a) Total Invasive Annual Cover (Grass plus Forb) 80 60 20 0 c) Native Annual Plant Diversity 12, a _ 09 = G6 2 = Ge 0 e) Native Annual Plant Richness Cover (%) Is O per Quadrat 3 a. p [oe b ee G2 0 |_|] Once-Burned Wi Twice-Burned b) Relative Cover of Invasive Annual Grass 80 60 b 40+ 4 20 0 d) Native Annual Plant Species Similarity 0.8 b pig ese we E04 0.2 0 f) Native Annual Plant Richness per Plot Cover (%) E25, a (a0) a 20 : 245 C g 10 a ° Gg o FIG. 3. Invasive annual plant cover (a), relative invasive annual grass cover (b), and various native annual plant diversity measures: Shannon index (c), Sorenson index (d), richness per 1 m?* quadrat (e), and richness per 168.3 m° mod-NAWMA plot (f). Values in each graph are averages per stand with SE bars. Differences in letters between paired stands within each graph indicate significant differences at « = 0.05. L] Unburned a) Invasive Grass Cover Cover (%) oO Cover (%) NO W a oO & sad) Oy = L) Once-Burned b) Invasive Forb Cover 60 60 50 : 50 : se 40 4o} ° = ab @ 30 30 6 20 a 20 10 10 0 0 M@ Twice-Burned c) Native Annual Cover 10 Fic. 4. Average cover and SE bars for invasive grasses (a), invasive forbs (b), and native annuals (c) in the unburned, once-burned, and twice-burned stands. Differences in letters above bars indicate significant differences between stands per graph based on ANOVA and LSD tests (« = 0.05). 20 MADRONO common following fire due to increased ash (Raison 1979; Abella et al. 2009). The response of total N and C to fire, however, is variable (Raison 1979; Brooks 2002; Allen et al. 2011). Pre-fire Encelia cover was assumed to be high in the twice-burned stand based on conditions in the once-burned stand, which likely accounts for the elevated C and N found in post fire soils (Xie and Steinberger 2001). Soil NHy* and NO; were highest in the twice-burned stand but not sig- nificantly so. These mobile, inorganic forms of nitrogen are typically elevated in post-fire envi- ronments (Wan et al. 2001), even in creosote bush scrub (Esque et al. 2010b). It is possible that both NH, and NO3 were significantly elevated im- mediately after the 2005 fire in the twice-burned stand but by the time the sites were sampled in August of 2006, these nitrogen sources had de- creased due to leaching and/or immobilization. Post-fire bare ground and rock cover was el- evated in the twice-burned stand. Adams et al. (1970) reported higher bare ground after fire in creosote bush scrub of the Colorado Desert due to hydrophobic soils that were characterized by water repellant layers found at various depths under burned shrubs. Large bare areas under burned shrubs, as they described, were not ob- served during this study. In general, the altered soil properties that resulted from fire were not dramatic and are expected to return to pre-fire conditions as vegetation recovers (Allen et al. 2011). However, persistent invasive species, a continuation of a short fire return interval or heightened soil erosion could cause long-term al- terations to soil properties (Morris and Moses 1987; Belnap 1995; Allen et al. 2011). Impact of Fire on Seed Bank Germinants In general, propagule abundance is linked to above ground plant performance (Olano et al. 2005; Cox and Allen 2008). Therefore, seed bank composition can elucidate potential aboveground vegetation, especially in the context of future disturbances (Cox and Allen 2008; Satterthwaite et al. 2007; Fisher et al. 2009). Results from the seed bank assays revealed that invasive grass propagules of Schismus spp. are ubiquitous and abundant in the seed bank at this site. Future fires or other disturbances will likely promote these plants (Cox and Allen 2008; Fisher et al. 2009). Native species also did not differ among stands, which suggests that invasive species re- moval could be an effective strategy for native seed bank management, especially because native annual plants exhibit density dependent inhibi- tion of germination (Inouye 1980). When scaled up, the number of invasive an- nual grass propagules in the twice-burned stand was 111,952.9 + 15,760 SE per m’*. To our knowledge, this is vastly greater than any value [Vol. 59 previously reported for exotic annual grasses from the American southwest (Young and Evans 1975; Nelson and Chew 1977; Reichman 1984; Hassan and West 1986; Guo et al. 1998; Angoa- Roman et al. 2005; Cox and Allen 2008; Abella et al. 2009; Esque et al. 2010a). The relative lack of other germinants besides Schismus spp. may indicate that the methods used to assay the seed bank were not ideal for detecting the full suite of species that could occur in the seed bank. Native desert annuals, in particular, are known to exhib- it high interannual variation (Freas and Kemp 1983; Philippi 1993; Pake and Venable 1996). While multiple watering cycles were utilized to address this potential variation, and while Schis- mus germinants were removed immediately to minimize interference, it is possible that not all viable native seeds in the seed bank samples ger- minated during the assays. For example, Esque et al. (2010a) treated seed bank samples with gib- berellic acid to stimulate germination and ob- served native annual germinants to be magni- tudes greater than what we found. Impact of Fire on Annual Plants Invasive annual grasses and forbs can severely reduce the abundance and species richness of native annual plants in unburned vegetation (Huenneke et al. 1990; Crimmins and McPherson 2008; Davies and Svejcar 2008; Minnich 2008). This study suggests that fire disturbance is also a serious threat to native annuals because it pro- motes invasive plants like Schismus spp. Abun- dance of Bromus madritensis ssp. rubens typically decreases in the immediate post-fire years (Abella et al. 2009; Esque et al. 2010a) although it 1s ex- pected to return to or even exceed pre-fire abun- dance levels within three years after fire, if pre- cipitation is adequate (Brooks 2003). The mechanism whereby Schismus spp. increases 1m- mediately after fires relates to the small size of its seeds, which fall into cracks and escape damage from fire, plus its ability to take advantage of elevated inorganic nitrogen levels in the post- fire environment (Esque et al. 2010b). Because nitrogen-use traits may not differ between in- vasive and native annuals, the relative early ger- mination and more rapid phenology of Schismus spp. contributes to its success (Marushia et al. 2010; Steers et al. 2011). At this study site, fire reduced the quadrat- and plot-level species richness of native annual plants, and recurrent fire magnified this outcome at the plot-level. Recurrent fire also significantly de- creased species diversity, which led to a highly simplified assemblage of annual plants in the twice-burned stand. Despite the negative impact of fire on native annuals, if invasive annuals are removed post-fire, then native annual species 2012] richness can increase greatly, likely exceeding pre- fire levels (Steers and Allen 2010, 201 Ic). Relevance to the Grass/Fire Cycle At our study site, invasive annual grass cover within the first three years after a fire was greater in the burned compared to the unburned stand, due almost entirely to Schismus spp, which 1s similar to other studies (Cave and Patten 1984; Minnich and Dezzani 1998: Brooks 2002; Esque et al. 2010b; Steers and Allen 20lla). This difference in invasive annual grasses may translate to greater potential for a consequent fire (Brooks et al. 2004). Schismus spp. are generally considered less effective at carrying fire than larger annual grasses like Bromus madritensis ssp. rubens (Brooks 1999). However, in this region Schismus spp. can attain relatively large sizes due to high anthropogenic nitrogen deposition, especially in wet years (Rao and Allen 2010; Rao et al. 2010) and are known to carry stand-replacing fires. For example, in the summer of 2005, at least four other creosote bush scrub fires within a 10 km radius of the study site were primarily fueled by Schismus spp. (R. Steers personal observations). Therefore, given adequate precipitation, results from this study suggest that fire can promote invasive annual grasses (1.e., Schismus spp.), which in turn, could fuel addi- tional fires in a positive feedback, as described by the grass/fire cycle (D’Antonio and Vitousek 1992; Brooks et al. 2004). Management Implications Removal of invasive grasses and forbs should favor natives through decreased competition (Brooks 2000; Schutzenhofer and Valone 2005; Barrows et al. 2009; Steers and Allen 2010) and through limiting future fire disturbance (Brooks et al. 2004). Because native annual richness is linked to the spatial and structural heterogeneity of creosote bush scrub (Schmida and Whittaker 1981), some native species may not find suitable micro-habitats until the shrub components are returned, regardless of invasive plant removal. For example, Pholistoma membranaceum was the most abundant annual forb in the unburned stand (Appendix |) where it occurred almost ex- clusively in shrub understories of long-lived spe- cies, like Larrea tridentata (R. Steers personal observation). Pholistoma membranaceum was vir- tually eliminated in the once-burned and twice- burned stands even though Encelia farinosa shrubs were prevalent. Unfortunately, reestab- lishment of long-lived shrubs, like Larrea triden- tata, has been speculated to take decades or longer (Vasek 1983; Lovich and Bainbridge 1999: Abella 2009, 2010; Steers and Allen 2011b). 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WRCC, Reno, NV. Website http://www. wrcc.dri.edu [accessed 7 June 2008]. XIE, G. AND Y. STEINBERGER. 2001. Temporal patterns of C and N under shrub canopy in a loessial soil desert ecosystem. Soil Biology and Biochemistry 33:1371-1379. YOUNG, J. A. AND R. A. EVANS. 1975. Germinability of seed reserves in a big sagebrush community. Weed Science 23:358—364. 24 MADRONO [Vol. 59 | APPENDIX 1. Average cover of each annual species recorded per quadrat (1 m’) and their respective frequency score per stand (unburned, once-burned, or twice-burned stands). Frequency is the number of occurrences per six 168.3 m° mod-NAWMA plots per stand. Unburned Once-burned Twice-burned | Cover (%)/ Cover (%)/ Cover (%)/ | frequency frequency frequency | Family Species per six plots per six plots per six plots | INVASIVE FORBS Brassicaceae Brassica tournefortii Gouan 22.1/6 41.9/6 5.2/6 Geraniaceae Erodium cicutarium (L.) Aiton 11.2/6 7.3/6 13.4/6 INVASIVE GRASSES Poaceae Bromus madritensis L. subsp. 4.1/6 <0.1/6 0/3 rubens (L.) Husn. B. tectorum L. 0/4 =< O.1y1 <0.1/1 Schismus barbatus (L.) Thell. 22.1/6 11.9/6 34.7/6 and S. arabicus Nees NATIVE ANNUALS Asteraceae Chaenactis fremontii A. Gray 3.1/6 0.8/6 0/2 Filago californica Nutt. 0.6/6 0/5 0.2/4 F. depressa A. Gray =—ULH/2 0/0 0/0 Lasthenia californica Lindl. <0.1/5 0/3 <0.1/5 Malacothrix glabrata A. Gray 0/0 0/3 0.2/3 Rafinesquia neomexicana A. Gray 0.1/6 0/0 0/0 Stephanomeria exigua Nutt. 0/0 0/0 = O1/2 Stylocline gnaphaloides Nutt. 1.4/6 O/1 0/0 Uropappus lindleyi (DC.) Nutt. 0/0 <0.1/1 0/1 Boraginaceae Amsinckia menziesii A. Nelson & 1/6 O/1 O/1 J. F. Macbr. Cryptantha barbigera Greene 0/5 0.8/6 1/5 Pectocarya heterocarpa 1. M. 0.3/5 0/0 0/0 Johnst. P. linearis DC. 0.7/5 <0.1/5 <0.1/1 P. recurvata 1. M. Johnst. 0.8/5 0.1/5 0/2 Brassicaceae Descurainia pinnata (Walter) 0/0 =0.173 O/1 Britton Lepidium lasiocarpum Torr. & A. 0/0 0/3 <0.1/1 Gray Tropidocarpum gracile Hook. O/1 0/0 0/0 Caryophyllaceae Loeflingia squarrosa Nutt. <0.1/5 0/0 0/0 Crassulaceae Crassula connata (Ruiz & Pav.) 1.3/6 0.1/4 <0.1/4 A. Berger Fabaceae Lotus strigosus (Nutt.) Greene =O1/5 0/0 =), 1/1 Lupinus sparsiflorus Benth. 0/2 0/0 0/0 Hydrophyllaceae Emmenanthe penduliflora Benth. 0.1/6 5.3/6 2.5/6 Phacelia campanularia A. Gray 0/0 0/2 =O. 1/2 P. distans Benth. 1.4/5 <0.1/6 0/0 Pholistoma membranaceum 13.2/6 O/1 0/0 (Benth.) Constance Lamiuaceae Salvia columbariae Benth. 0/0 0/0 0/1 Loasaceae Mentzelia involucrata S. Watson 0/0 O/1 0/0 Mentzelia sp. =—Uy2 0/0 0/0 Onagraceae Camissonia californica (Torr. & 0.1/6 1.1/6 2/6 A. Gray) P. H. Raven C. pallida (Abrams) P. H. Raven 0.1/4 0/2 <0.1/4 Plantaginaceae Plantago ovata Forssk. 0.1/6 0.7/6 1.11/3 Poaceae Vulpia microstachys (Nutt.) Benth. O/1 0/0 0/0 V. octoflora (Walter) Rydb. 3.9/6 0.9/6 0/1 Polemoniaceae Gilia angelensis V. E. Grant 0.2/1 0/0 0/0 Linanthus bigelovii Greene 0/2 0/0 0/0 Polygonaceae Chorizanthe brevicornu Torr. 0/2 0/0 0/0 Pterostegia drymarioidesFisch. 0/1 0/0 0/0 & C. A. Mey. Portulaceae Calyptridium monandrum Nutt. 0/0 0/1 O/1 MADRONO, Vol. 59, No. 1, pp. 25-27, 2012 STATUS OF BINGHAM’S MORNING-GLORY IN THE LIGHT OF ITS REDISCOVERY R. K. BRUMMITT The Herbarium, Royal Botanic Gardens, Kew, Richmond Surrey, TW9 3AE, U.K. ScoTT D. WHITE! AND JUSTIN M. Woop Aspen Environmental Group, 201 North First Ave., No. 102, Upland, CA 91786 swhite@aspeneg.com ABSTRACT Calystegia sepium (L.) R. Br subsp. binghamiae (Greene) Brummitt (Convolvulaceae), until recently presumed extinct, is elevated to species status. The basionym Convolvulus binghamiae Greene was published without identifying a type; therefore, a lectotype is selected from among the specimens cited in Greene’s description. Key Words: Calystegia sepium subsp. binghamiae, Convolvulaceae, lectotype, new combination, rare species. Calystegia sepium (L.) R. Br subsp. binghamiae (Greene) Brummitt has been presumed extinct (California Native Plant Society 2011) until its rediscovery in May 2011 in the City of Chino, San Bernardino County, California. The redis- covery and subsequent conservation efforts will be described elsewhere by others. The availability of new specimens and live material prompted a taxonomic review, which indicates that recogni- tion at the species level is warranted. TAXONOMIC TREATMENT Calystegia binghamiae (Greene) Brummitt, comb. nov.—Basionym: Convolvulus binghamiae Greene, Bull. Calif. Acad. Sci. 2: 417. 1887. Greene referred to collections by Bingham and himself “‘in marshy places about Burton’s Mound in Santa Barbara,” but did not cite the specimens. Synonyms: Convolvulus sepium var. binghamiae (Greene) Jepson, Fl. Calif. 3:118. 1939. Calystegia sepium subsp. bingha- miae (Greene) Brummitt, Ann. Missouri Bot. Gard. 216. 1965.—Type: USA, California, Santa Barbara Co. City of Santa Barbara, August 1886, Mrs. R.F. Bingham s.n. UC 335392 (lectotype chosen here; isolectoype: Mrs. R.F. Bingham s.n., Columbian Collection F). Jepson cited Bingham’s collection, but did not specify the UC or the F specimen as the lectotype. Review Convolvulus binghamiae Greene (Convolvula- ceae) was described from specimens collected by Mrs. R. F. Bingham and E. L. Greene in Santa ' Author for correspondence Barbara, coastal southern California, in 1886. In 1965 Brummitt transferred it to the genus Calystegia R. Br. and ranked it as a subspecies within C. sepium (L.) R. Br., a decision he has regretted since. Its only verified localities are coastal regions of Santa Barbara and Los Angeles counties and Chino Creek, San Bernar- dino County, all in southern California (Consor- tium of California Herbaria 2011). Abrams (1951) also mentioned it extending to Orange County. We have seen specimens from Bolsa Chica (L.M. Booth 1214, POM) and east of Huntington Beach (L.M. Booth 1359, POM) that were labeled as Convolvulus binghamiae or C. sepium subsp. binghamiae as of 1951. Both of these were annotated by Brummitt as Calystegia sepium subsp. limnophila (Greene) Brummitt. We are not aware of any other records from Orange County. One of us, Brummitt, has worked on this genus for many years, both in the herbarium and in the field. In 1998 he determined material at RSA as Calystegia binghamiae, adopting specific rank, but, in view of the lack of clear evidence in the very sparse material available to him, he did not publish this name. In The Jepson Manual (Brummitt 1993) and its second edition (Baldwin et al. 2012), which went to press before the rediscovery was appreciated, he retained subspe- cific rank under C. sepium. However, he has now examined the recent specimen collected in Chino (J.M. Wood 4092, K), as well as Greene’s original description, and photographs of the original material, and as a result is now convinced that inclusion in C. sepium is inappropriate. In C. sepium, with numerous subspecies in pan- temperate regions of the world, the large paired bracteoles are inserted close to the calyx and largely overlap and conceal it. This seems to be 26 MADRONO an apomorphy suggesting a derived position in the likely evolution of the genus. Calystegia binghamiae, by contrast, usually (see below) has smaller, much narrower bracteoles, with at least one of them inserted remote from the calyx. Such bracteoles are characteristic of a number of species of the Calystegia complex that is endemic to California. This character is not found elsewhere among Ca/ystegia taxa, and 1s thought to represent a more plesiomorphic condition. Calystegia sepium may well be a polyphyletic taxon (Brummitt 1963) even without including C. binghamiae, and would probably be even more so with C. binghamiae included (the taxonomic details of the Californian species in the 1963 thesis were based on inadequate herbarium material and have been superseded by the author in more recent work). The rhizomatous habit, which C. binghamiae shares with C. sepium, apparently evolved independently within the California Calystegia lineage. Although available specimens referable to C. binghamiae are rather limited in number, we have noted surprising variation in both bracteole and leaf shape characters. The bracteoles on the original collections by Bingham and Greene from Santa Barbara are broadly elliptic, 8-12 x 4— 8 mm, and inserted almost adjacent to the sepals (probably influencing earlier decisions to include the taxon in C. sepium). They differ markedly, however, from those of C. sepium in being only about half as long as the sepals. All specimens we have seen from further east have linear to narrowly elliptic bracteoles with at least one inserted clearly below the sepals. Field observa- tions at different times by one of us (J. M. Wood) on the newly located population at Chino have noted that early in the season the bracteoles tend to be more similar to those of the specimens collected by Bingham and Greene in Santa Barbara, whereas later in the season they are much narrower with at least one of them usually remote from the sepals. A good illustration of the latter may be seen in Abrams (e.g., Fig. 3855, 1951); The leaves on the material from Santa Barbara have relatively well developed posteriorly-direct- ed basal lobes with a tendency to a parallel-sided sinus. Ivan Johnston’s specimens, collected at Chino Creek in 1917 (J274, below) have very similar leaves. However, the new collection from Chino, J.M. Wood 4092, has poorly developed basal lobes (especially on young leaves) with a broadly rounded sinus or almost cuneate leaf base. This is unlike anything found in C. sepium. Further specimens or observations on both leaves and bracteoles would be of interest. While excluding Calystegia binghamiae from C. sepium, one must consider whether it is possible to regard it as conspecific with any other Californian species, but this does not seem to be [Vol. 59 the case. Indeed it is not clear which of the Californian taxa would be most similar based on character states of the rhizomes, pubescence, leaves and bracteoles. An annotation made by Brummitt in 1973 on one the Johnston specimens at RSA suggested it was a hybrid possibly between C. sepium subsp. limnophila and C. occidentalis (Gray) Brummitt subsp. fulcrata (Gray) Brummitt or C. /ongipes (Watson) Brum- mitt, but this is now discounted. One misidentified specimen, R. Zembal s.n. 21 May 1977 RSA, labeled as Calystegia sepium subsp. binghamiae and reported as such in California Department of Fish and Game (2011), is C. macrostegia (Greene) Brummitt. One of us (S. D. White) has annotated the specimen and entered the correction on the Consortium of California Herbaria web site. Calystegia binghamiae has been known by the common names “Santa Barbara morning-glory”’ (Abrams 1951) and ““Bingham’s false-bindweed”’ (USDI Natural Resources Conservation Service 2011). Its geographic range is (or was) wider than the first common name implies, and the native California Calystegia species are commonly known as morning-glories rather than false- bindweeds (Brummitt 1993). Therefore, we sug- gest the common name “Bingham’s morning- glory.” Mrs. R.F. Bingham was a naturalist of the Santa Barbara area, and published notes on the local vascular flora, marine algae, natural history, and medicinal plants (e.g., Bingham 1887, 1890). A genus of marine algae, Bingha- miella, is named in honor of a Mrs. C. P. Bingham of the Santa Barbara area in the 1870s (Setchell and Dawson 1941); this may have been the same Mrs. Bingham, perhaps identifying herself at times by her husband’s initials. Specimens Examined USA. CALIFORNIA. Santa Barbara Co.: Santa Barbara, August 1886, Mrs. R.F. Bingham sn. (UC 335392, lectotype chosen, here; FP, presumed duplicate of previous cited collection; photos K, RSA); Santa Barbara, 1886 E.L. Greene s.n. NDG 39692, and July 1886, E.L. Greene s.n. NDG 39691 and 39693; photos K, RSA); lagoon near ocean, Ellwood, May 30 [no year] Alice Eastwood s.n. (UC 879470). Los Angeles Co.: Riveria [probably what is now Pico Rivera], | May 1902, Anstruther Davidson 1892 (RSA; mixed collection with one stem of C. binghamiae including leaves, one flower, and one bud, mounted with several C. sepium stems); near University Station [presumably a Pacific Electric station near the USC campus], 1899, Anstruther Davidson 2144 (RSA). San Bernardino Co.: Chino Creek, 30 May 1917, Ivan Johnston 1274 (two sheets at RSA/POM:; one at UC); city of Chino, SE corner of Edison Ave. and Oaks Ave., near 2012] entrance to Chaffey college campus, ca. 2.5 mi N of Chino Creek (Prado Basin), irrigated land- scaped area adjacent to ruderal grasslands, 17 May 2011, Justin M. Wood 4090 (to be distrib- uted) and 4092 with S.D. White, N. Gale & A. Parikh (K, RSA; one duplicate to be distributed). We understand that at least one other collection has been made at the Chino site this year, but we have not seen it. ACKNOWLEDGMENTS We thank Mitchell C. Provance, Andrew C. Sanders, and two anonymous reviewers for their assistance in preparing this note. LITERATURE CITED ABRAMS, L. R. 1951. Llustrated flora of the Pacific states, Vol. 3. Stanford University Press, Stanford, CA. BALDWIN, B. G., D. H. GOLDMAN, D. J. KEIL, R. PATTERSON, AND T. J. ROSATTI (eds.). 2012. The Jepson manual: vascular plants of California. 2nd ed. University of California Press, Berkeley, CA. BINGHAM, R. F. 1887. Flora near Santa Barbara, California. Botanical Gazette 12:33—35. BRUMMITT ET AL.: BINGHAM’S MORNING-GLORY ZF . 1890. Medicinal plants growing wild in Santa Barbara and vicinity. Bulletin of the Santa Barbara Society of Natural History 1:34—37. BRUMMITT, R. K. 1963. A taxonomic revision of the genus Calystegia. Ph.D. thesis, University of Liver- pool, UK. . 1993. Calystegia. Pp. 517-521 in J. C. Hickman (ed.), The Jepson manual: higher plants of Califor- nia. University of California Press, Berkeley, CA. CALIFORNIA DEPARTMENT OF FISH & GAME. 2011. California Natural Diversity Database Version 3.1.1. Heritage Section, CDFG, Sacramento, CA. CALIFORNIA NATIVE PLANT SOCIETY. 2011. Inventory of rare, threatened, and endangered plants of California, v8-Ola. Sacramento, CA, Website: http://www.rareplants.cnps.org/ [accessed 12 July 2011]. CONSORTIUM OF CALIFORNIA HERBARIA. 2011. Web- site: http://ucjeps.berkeley.edu/cgi-bin/get_consort. pl [accessed 12 July 2011]. SETCHELL, W. A. AND E. Y. DAwson. 1941. Bingha- mia, the alga, versus Binghamia, the cactus. Proceedings of the National Academy of Science (USA) 27:376—-381. USDI NATURAL RESOURCES CONSERVATION SERVICE. 2011. PLANTS Database. http://plants.usda.gov/ java/ [accessed 16 September 2011]. MADRONO, Vol. 59, No. 1, p. 28, 2012 CHANGE IN RANK OF ERIODICTYON TRASKIAE SUBSP. SMITHIT (HY DROPHY LLACEAE) GARY L. HANNAN Department of Biology, Eastern Michigan University, Ypsilanti, MI 48197 ghannan@emich.edu In preparing a taxonomic treatment of Erio- dictyon (Hydrophyllaceae) for the Flora of North America North of Mexico, it has become necessary to standardize ranks of infraspecific taxa. Infraspecific taxa in two species in the Flora have been named as varieties: E. trichocalyx A. Heller var. trichocalyx with E. trichocalyx var. lanatum (Brand) Jeps., and, E. crassifolium Benth. var. crassifolium with E. crassifolium var. nigrescens Brand. The operationally, if not evolutionarily, preferable approach to achieving nomenclatural consistency in the treatment is to treat E. traskiae Eastw. subsp. smithii Munz as a variety. In the interest of nomenclatural consis- tency, therefore, the following nomenclatural change is proposed: TAXONOMIC TREATMENT Eriodictyon traskiae Eastwood var. smithii (Munz) Hannan stat. nov. Basionym: Eriodictyon tras- kiae Eastw. subsp. smithii Munz, A California Flora: Supplement, 90. 1968. —Type: USA, California, Santa Barbara Co., San Marcos Pass, July 4, 1950, Clifton F. Smith 1621 (holotype: POM 310992; isotypes: CAS 384681!, NY 337110). The resulting autonym 1s: Eriodictyon traskiae Eastw. var. traskiae —TY PE: USA, California, Los Angeles Co., Santa Catalina Island, Avalon, “‘one volcanic upland, 1500 ft. elevation’, May, 1897, Blanche Trask s.n. (holotype: CAS 369!). MADRONO, Vol. 59, No. 1, pp. 29-43, 2012 MIMULUS SOOKENSIS (PHRYMACEAE), A NEW ALLOTETRAPLOID SPECIES DERIVED FROM MIMULUS GUTTATUS AND MIMULUS NASUTUS BEVERLY G. BENEDICT Connell Herbarium, University of New Brunswick, 10 Bailey Drive, Fredericton, N.B., Canada E3B 5A3 JENNIFER L. MODLISZEWSKI Department of Biology, Duke University, Campus Box 90338, Durham, NC 27708 jlmS50@duke.edu ANDREA L. SWEIGART Department of Genetics, University of Georgia, Fred C. Davison Life Sciences Complex, Athens GA 30602 NOLAND H. MARTIN Department of Biology, Texas State University-San Marcos, 601 University Drive, San Marcos, TX 78666 FRED R. GANDERS Department of Botany, University of British Columbia, 3529-6270 University Blvd. Vancouver B.C., Canada V6T 124 JOHN H. WILLIS Department of Biology, Duke University, Campus Box 90338, Durham NC 27708 ABSTRACT A new species of monkeyflower, Mimulus sookensis, is described. This species is found throughout the southern portion of Vancouver Island, the Gulf Islands of British Columbia, the San Juan Islands of Washington state, the Willamette and Umpqua River Valleys in Oregon, and has been collected at one location in Mendocino County, California. Mimulus sookensis is a tetraploid species (7 = 28) derived from the predominately outcrossing Mimulus guttatus DC. (n = 14) and the predominately self-pollinating Mimulus nasutus Greene (n = 14). Mimulus sookensis is similar phenotypically to the small-flowered M. nasutus, but differs in chromosome number, height, and by a slightly more narrowed corolla tube than that of M. nasutus. It is commonly found on wet hillsides, seeps, cutbanks, and in roadside ditches, often co-occurring with M. guttatus but infrequently with M. nasutus. Key Words: Allotetraploid speciation, Mimulus, Mimulus guttatus, Mimulus nasutus, monkeyflower, new species, Oregon, Vancouver Island. A small-flowered monkeyflower similar to Mimulus nasutus Greene was first observed on Vancouver Island, Canada, by Fred Ganders, and later collected for scientific study in May 1991 by Beverly Benedict. Although phenotypically simi- lar to M. nasutus (Fig. 1), allozyme analysis revealed that some of the small-flowered monkey- flowers on Vancouver Island were always hetero- zygous at allozyme markers. This was in contrast to allozyme data from another small-flowered monkeyflower found on the island, M. nasutus (snouted monkeyflower), and the common yellow monkeyflower, M. guttatus DC. These results were intriguing because while the large-flowered, chasmogamous M. guttatus is known to be highly outcrossing, both M. nasutus and the ‘Author for correspondence heterozygous, small-flowered monkeyflowers were known to be highly selfing, given their floral structure, small flower size, and often cleistoga- mous nature (Ritland and Ritland 1989; Dole 1992; Willis 1993). Morphological analysis of M. guttatus, and the two small-flowered monkey- flowers (M. nasutus and the species described here, M. sookensis) revealed that while WM. nasutus and M. sookensis overlapped a great deal in floral morphology, subtle morphological differences did exist (Fig. 1, e.g., pistil length, corolla tube width). Because of fixed heterozygosity in some of the small-flowered Mimulus on Vancouver Island, and slight differences in floral morphology, F. Ganders suspected that the heterozygous mon- keyflowers in question were actually a distinct taxon of allopolyploid origin (Benedict 1993). Chromosome squashes conducted at the time revealed that these new monkeyflowers, M. 30 MADRONO Fic. 1. [Vol. 59 fe. ts “wo Photographs of M. sookensis and its progenitor species. Side profile photographs are taken on approximately the same scale. A. M. sookensis, B. M. guttatus, C. M. sookensis, D. M. nasutus. sookensis, had more than n = 14 chromosomes, but an exact count was not obtained. Following the work of B. G. Benedict, flow cytometry data from three M. sookensis collec- tions revealed that the small-flowered monkey- flowers from Vancouver Island and surrounding areas, as well as the valleys of western Oregon and northern California, had approximatey twice the DNA content of M. guttatus and M. nasutus, suggesting again that this taxon was of tetraploid origin (Sweigart et al. 2008). Sequence data from two nuclear genes confirmed that this new species was a hybrid tetraploid derived from M. guttatus and M. nasutus. Furthermore, crossing data revealed that the allotetraploids were reproduc- tively isolated from their diploid progenitors due to failure of seed development, a result consistent with the triploid block that is commonly ob- served in interploidy crosses (Sweigart et al. 2008). Although M. sookensis is a cryptic species due to its phenotypic similarity to M. nasutus, the fact that it is reproductively isolated from its diploid progenitors illustrates the concept of instant or rapid speciation of polyploids, which has long been recognized (e.g., Winge 1917; Dobzhansky 1937; Coyne and Orr 2004). Poly- ploidy not only has the propensity to quickly create new species (according to the biological species concept, e.g., Mayr 1996) but has contributed significantly to angiosperm evolution (Stebbins 1971; Grant 1981; Masterson 1994; Otto and Whitton 2000). Here, we present evidence that M. sookensis is historically taxonomically unrecognized, and provide new chromosome data that provide conclusive evidence that M. sookensis is a cytologically distinct species, which has previous- ly been shown (Sweigart et al. 2008) to be of polyploid origin, and reproductively isolated from its diploid progenitors, as well as a description of this hitherto unnamed species of monkeyflower. REVIEW OF PREVIOUSLY PUBLISHED MIMULUS TAXA Mimulus guttatus is an herbaceous wildflower distributed throughout much of western North America (Vickery 1978), while Mimulus nasutus has a restricted range relative to M. guttatus (Kiang and Hamrick 1978; Vickery 1978). Mimulus guttatus, M. nasutus, and M. sookensis all belong to the M. guttatus species complex, and are part of the Simiolus clade (Beardsley et al. 2004) of the genus Mimulus. Mimulus guttatus and its close relatives have been extensively 2012] BENEDICT ET AL.: SHY MONKEYFLOWER-A NEW POLYPLOID MIMULUS 3] TABLE 1. COMPLETE LIST OF PREVIOUSLY PUBLISHED MIMULUS TAXA WHICH MIGHT HAVE BEEN A DESCRIPTION OF M. SOOKENSIS, WITH A DESCRIPTION OF HOW THEY ARE DIFFERENT FROM M. SOOKENSIS. Taxa are listed in alphabetical order, although subspecies and varieties are listed in parentheses if variety or subspecies was given specific rank. For synonyms examined, three sources were used: the synonyms listed in Grant (1924) and Pennell (1951) for M. nasutus, and the synonyms listed for both M. guttatus and M. nasutus in IPNI. Many of the large flowered varieties of M. guttatus were not included in this list for the purpose of brevity. All references are included in the literature cited. Evidence sources refers to all herbarium specimens, drawings and descriptions, in both the nomenclatural citation and established floras or monographs, that were used in determining differences. For each candidate taxa, the characters that most easily illustrate the difference between the listed taxa and M. sookensis are described for the listed taxa. Previously published taxa Synonym (source) Evidence sources Distinguishing characters from M. sookensis M. arvensis Greene (M. guttatus DC. var. arvensis Grant) M. bakeri Gandoger M. cordatus Greene M. cuspidatus Greene M. decorus (Grant) Suksd. (M. guttatus DC. var. decorus Grant) M. erosus Greene M. glareosus Greene M. guttatus DC. subsp. scouleri (Hook.) Pennell M. hallii Greene (M. guttatus DC. var. hallii Grant) M. guttatus DC. var. /yratus (Benth.) Pennell ex M. Peck M. guttatus var. depauperatus Grant (M. luteus var. depauperatus A. Gray) M. guttatus var. glaucescens (Greene) Jeps. (M. glaucescens) M. guttatus (IPNI) M. nasutus (Grant 1924, Pennell 1951) M. nasutus (Grant 1924) M. nasutus (Grant 1924; Pennell 1951) M. guttatus (IPNI) M. nasutus (Grant 1924) M. nasutus (Grant 1924; Pennell 1951) M. guttatus (IPNI) M. guttatus (IPNI) M. guttatus (IPNI) M. guttatus (IPNI) M. guttatus (IPNI) Greene (1887); Grant (1924): Pennell (1951); Mukherjee and Vickery (1962) CAS 22488 (isotype), NY 20798 (possible isotype); Gandoger (1919) Greene (1910); Pennell (1951); Mukherjee and Vickery (1962) DS 771002 (isotype); description in Greene (1910) CAS 22445; Pennell (1951) Greene (1910) Greene (1889, 1894) Pennell (1947) Greene (1885); Grant (1924) Pennell (1941); Pennell (1951) Gray (1867); Grant (1924): Hitchcock and Cronquist (1987) Greene (1885); Jepson (1925); Pennell (1951) Diploid (7 = 14), easily hybridizes with M. guttatus; Greene describes the leaves as lyrate, and the species as perhaps synonymous with M. /yratus Benth. Grant describes the variety as having an upper calyx tooth not markedly longer than others, elongated internodes, teeth not usually folded over each other at maturity CAS specimen appears to be hybrid between M. guttatus and M. nasutus, while description doesn’t match specimen, description suggests that difference between M. nasutus and M. bakeri is the impunctate calyx of M. bakeri Corolla lacking in spotting, diploid (7 = 14) that hybridizes easily with M. guttatus M. nasutus found in wet shades exhibiting phenotypic plasticity in a classic shade avoidance response (see text for discussion) Corolla large Corolla exserted from tube, synonym of M. nasutus Leaves toothed or lobed, slimy, synonym of M. nasutus Stoloniferous variant of M. guttatus with more linear leaves (perhaps synonymous with M. filingii Regel or M. caespitosus Greene) Leaves parallel-veined and almost entire, calyx highly inflated Leaves pinnately lobed at the base, corolla long (2—3 cm) Grant (1924) thought to be synonymous with M. puncticalyx and M. microphyllus, based on Hitchcock and Cronquist (1987) and Gray, appears to be simply a description of small M. guttatus or M. nasutus plants with few or small flowers — a condition most likely caused by environment Leaves glaucous, synonymous with M. g/aucescens (Greene) 32 MADRONO [Vol. 59 TABLE 1. CONTINUED. Previously published taxa Synonym (source) Evidence sources CAS 23523 (isotype for Distinguishing characters from M. sookensis M. guttatus var. gracilis (A. Gray M. guttatus (IPNI) Campbell lumps all synonyms of M. nasutus and M. nasutus itself under this variety. CAS specimen is M. ex Torr.) M. pardalis); pardalis, corolla described as being twice as long as Campbell Campbell the calyx, diploid (n = 14) (1950); ORE96554 M. nasutus (Pennell 1951) M. guttatus M. guttatus var. nasutus Jeps. M. guttatus var. Synonym of M. nasutus Grant (1924) Listed as perennial, large-flowered puberulus A. L. (IPNI) Grant M. inflatulus M. breviflorus CAS 152750 Calyx equal-toothed, leaves more linear and narrow, Suksd. (Pennell 1951) (isolectotype); synonym of M. breviflorus Piper Pennell (1951) M. laxus Pennell M. guttatus CAS 329746 Variant of M. guttatus, diploid (n = 14) ex. M. Peck (Mukherjee and Vickery 1962) (isotype); NY 90734 (isotype); Mukherjee and Vickery (1962) M. nasutus Greene (1895b) (Pennell 1951) M. marmoratus Greene Description of M. marmoratus matches that of a hybrid between M. guttatus and M. nasutus, with large red blotch on middle lower lobe, with a corolla that is longer than M. nasutus (>3 cm) Narrow-range endemic of CA; calyx even toothed and lower teeth not curled upward and inward upon maturity, stem weak, lower leaves described as being lyrate and long-petioled, calyx puberulent, diploid used in multiple genetic studies (see text) DS 74105; NY 90746 (isotype Heller 7410); Heller (1912); Grant (1924): Pennell (1951); Munz (1959) M. micranthus A. Heller (MM. guttatus var. micranthus (A. Heller) G. R. Campb., M. nasutus Greene var. micranthus M. guttatus, M. nasutus (IPN1I) A. L. Grant) M. microphyllus Greene (1885), Leaves small, stems rounded, pistil much exserted from Benth. (M. Pennell (1941, calyx, located mostly in the mountains guttatus var. microphyllus Pennell in M. Peck) M. minusculus Greene M. minutiflorus R. K. Vickery M. nasutus Greene var. exiImius Green A. L. Grant ex J. T. Howell M. nasutus Greene var. insignis A. L. Grant M. guttatus DC. var. insignis Greene M. parishii Gand. M. puberulus Greene M. puberulus Gand. M. nasutus (Grant 1924) M. nasutus (IPNI) M. nasutus (IPNI) M. guttatus (IPNI) M. nasutus (Grant 1924: Pennell 1951) M. nasutus (Grant 1924: Pennell 1951) 1951) Greene (1910) CAS 961575 (isotype); Vickery (1997) Howell (1949) Grant (1924): and Pennell (1941) JEPS 2938 (the very type!) Gandoger (1919) Greene (1910) Gandoger (1919) Perennial, shorter than M. sookensis, leaves ovate, flowers large Corolla superficially similar in appearance to M. sookensis, but lacking ridges, and stems wiry; closely related to M. wiensii, n = 32 Howell (1949) bases his description of this variety on M. nasutus, but does not realize that what he considers M. nasutus is actually a hybrid between M. guttatus and M. nasutus, also appears to be synonymous with M. nasutus var. insignis Flower size outside the range of M. sookensis and large blotch of anthocyanin spotting on lower corolla lobe, both suggest that description matches that ofa hybrid between M. guttatus and M. nasutus Large flowered, hybrid between M. guttatus and M. nasutus Leaves deeply cut or laciniate; only a single specimen was examined in the naming Corolla large (>3 cm), stem round and viscidly puberulent Only distinguishing feature from typical M. nasutus is that it is minutely pubescent; only a single specimen was examined, a synonym of M. nasutus 2012] TABLE 1. Previously published taxa Synonym (source) M. puncticalyx M. nasutus ORE96654 Gand. (Pennell 1951) (isotype); ORE96655; Gandoger (1919) M. subreniformis — M. nasutus UC 2711 Greene (Grant 1924; (holotype); Pennell 1951) M. washingtonensis CAS 152669 Gand. (isotype); Gandoger (1919) collected and examined throughout western North America, by both early botanists and contemporary botanists and geneticists. Histori- cally, M. guttatus and its close relatives have been subject to extraordinarily divergent taxonomic treatments by different authors. Pennell (1951) recognized 28 taxa closely allied with M. guttatus from the Pacific Northwest, and in a recent treatment of California, Thompson (1993) recog- nized only five. In contemporary times, the genus Mimulus has seen a proliferation of scientific interest: a Google Scholar search for articles published between 1980-2011 including the word Mimulus in the title found 436 articles, with 194 written on M. guttatus alone. Although many of these recent publications do not necessarily include field work, it is safe to assert that more has been learned of the genetics, ecology, distribution, and taxonomic status of M. guttatus and its close relatives, since the publications of Grant (1924), Pennell (1951) and even Thompson (1993), see Wu et al. (2008). By combining knowledge from contemporary studies with historical taxonomic wisdom, we found that M. sookensis is truly a previously overlooked species in this intensely studied group, in part due to its cryptic nature. To determine if M. sookensis was previously taxonomically recognized, we first identified synonyms of M. guttatus (only the small-flowered or obscure taxa) and M. nasutus, from those listed in Pennell (1951), Grant (1924), and Campbell (1950), and from lists of synonyms derived from IPNI (International Plant Names Index). We also searched in Pennell (1951) and Grant (1924) for descriptions of small, yellow- flower Mimulus that were not listed as synonyms of M. guttatus or M. nasutus, but were considered to be closely related to the Simiolus clade (candidate taxa, Table 1). For these 31 candidate taxa, in which the author might have potentially described M. sookensis, we referred to herbarium specimens, the original species descriptions, crossing data and chromosome counts (when Evidence sources Greene (1895a) BENEDICT ET AL.: SHY MONKEYFLOWER-A NEW POLYPLOID MIMULUS a3 CONTINUED. Distinguishing characters from M. sookensis Leaves tiny upper tooth hardly more prominent than others; only a single specimen was examined in the naming Appears to be a diminuitive variant of M. nasutus, but without anthocyanin spotting on corolla Calyx equal-toothed, flowers large available), and drawings and descriptions in other references to determine if a_ previously published name could be applied to M. sookensis (Table 1). We did not find a previously published taxon that satisfied every aspect of the morphol- ogy and cytology of M. sookensis (Table 1), and thus, despite the abundance of synonyms within the M. guttatus species complex, no previously published names can be applied to M. sookensis. Throughout the course of our examination of M. sookensis candidates, we found that the reasons why candidate taxa were not representa- tive of M. sookensis fell into one or more categories. First, pronounced differences in habit, leaf, and even floral morphology existed (e.g., perenniality, lyrate leaves, even-toothed calyx). Second, in some cases the species described was likely either a hybrid between M. guttatus and M. nasutus, or M. nasutus. In the field, M. guttatus and M. nasutus are known to hybridize when they co-occur (Kiang 1973; Martin and Willis 2007). Hybrids between M. guttatus and M. nasutus have flowers that are much more similar in size to M. guttatus, due to dominance of the M. guttatus floral genes (Fishman et al. 2002). In the field, a prominent red blotch has often been observed on the lower middle corolla lobe of both M. nasutus (e.g., Pennell 1951; Kiang 1973) and some monkeyflowers with larger flowers than those of typical M. nasutus, but bearing resemblance to M. nasutus in shoot architecture and _ leaf morphology. This prominent red blotch has not been observed on M. sookensis flowers. The fact that the species described often had both larger flowers and a large red blotch suggests that they are either M. nasutus or hybrids between ™. guttatus and M. nasutus. Third, there were some cases in which floral morphology differences were subtle, but differences in chromosome number existed, based on crossing studies and chromo- some counts of Vickery (Campbell 1950; Mu- kherjee and Vickery 1962). In the special case of Mimulus micranthus A. Heller, it is defined in part by its endemism (Munz 1959). Mimulus 34 MADRONO TABLE 2. [Vol. 59 LIST OF COLLECTIONS USED IN MEIOTIC CHROMOSOME COUNTS AND IN THE PREVIOUSLY PUBLISHED FLOW CYTOMETRY ANALYSES PRESENTED IN SWEIGART ET AL. (2008). Abbreviations: MCC, meiotic chromosome count; FC, flow cyometry. Collection Taxon Locale Longitude Latitude Analyses DRN (DEX) M. sookensis Dexter’s Reservoir, OR, USA —122.756 43.917 FC LSN M. sookensis Lowell, OR, USA — 122.784 43.930 MCC, FC NHI M. sookensis Nanoose Hill, VI, BC, CAN — 124.160 49.273 MCC ROG M. sookensis ca. 12 mi SE of Marial, (as the —123.644 42.657 MCC, FC crow flies) OR, USA TRT M. nasutus near Troutdale, OR, USA — 122.368 45.520 MCC micranthus is a diploid that has been used in multiple genetic analyses, and has been success- fully crossed with other known diploids (Fenster and Ritland 1992, 1994: Ritland et al. 1993; Fenster et al. 1995). Last, we believe that the species described in some cases were possibly representative of phenotypic plasticity, the most noteable being M. cuspidatus Greene, found growing in shaded spots, with elongated inter- nodes, and lack of anthocyanin spotting. In Impatiens capensis Meerb., this phenotype 1s known to be an adaptive plastic response (Schmitt et al. 1995; Dixon et al. 2001) that is characteristic of the classic shade avoidance syndrome (Smith 1982; Smith and Whitelam 1997). While it is not possible to directly test for plasticity 1n previously collected specimens, it seems highly plausible that many of the candidate taxa that we examined are representative of either phenotypic variation or plasticity in M. nasutus or M. guttatus. Grant (1924) noted that ™. nasutus appeared to be quite a plastic species, and thus the taxa’s earlier designations (e.g., Grant 1924; Pennell 1951) as synonyms are appropriate. Additionally, Kiang (1973) demonstrated that Mimulus nasutus is an exceptionally plastic species, as the flower size is dependent upon both external environmental conditions, and the posi- tion of the flower along the stem. It is also well known that M. guttatus harbors a great deal of phenotypic variation (reviewed in Wu et al. 2008). CYTOLOGICAL ANALYSIS Meiotic counts of chromosomes were conduct- ed to corroborate the previous indications of polyploidy as evidenced by flow cytometry (Table 2), crossing barriers, (Sweigart et al. 2008), and fixed heterozygosity at allozymes and sequenced nuclear loci (Benedict 1993; Sweigart et al. 2008). Three individuals, each from different collections considered to be M. sookensis (Table 2, LSN, NHI, ROG) were used for the chromosome counts. A single diploid M. nasutus individual (TRT) was also counted, for the purpose of comparing chromosome sizes. Immature flower buds were collected in a 3:1 95% ethanol:glacial acetic acid solution. The tissue was transferred to 70% ethanol after 24 hr and stored at —20°C until ready for use. Flower buds were then partially dissected in a 70% ethanol solution. The partially-dissected floral material was then transferred to a half-strength aceto- carmine solution, where all non-anther material was removed. Anthers were then transferred to a drop of aceto-carmine on a slide, and were eviscerated to release the pollen mother cells from the anthers. After thorough evisceration, the tissue was removed from the solution, and the slide was placed on a warming plate to facilitate staining. A drop of Hoyer’s solution (Anderson 1954) was then added and the chromosomes were squashed by placing a coverslip over the solution and pressing down. Stained cells were examined with brightfield microscopy at 630-1000 mag- nification using a Zeiss Axioplan 2 microscope, and photographed at 1000 with a mounted Axiovision HR camera. Meiotic chromosome counts revealed 28 dis- tinct chromosome pairs in M. sookensis and 14 distinct chromosome pairs in diploid M. nasutus (Fig. 2). Although the sister chromatids are not easily distinguishable, it is clear from the chromosome squashes that there are twice as many of the chromosomes in M. sookensis as there are in diploid M. nasutus. This chromosome count constitutes the first published count for M. sookensis. Using these chromosome numbers as a calibration, we were able to confirm that the specimens used in the flow cytometry analysis of Sweigart et al (2008, Table 2) were indeed allotetraploid. TAXONOMIC TREATMENT Mimulus sookensis B. G. Benedict, J. L. Modlis- zewski, A. L. Sweigart, N. H. Martin, F. R. Ganders, and J. H. Willis, sp. nov. —TYPE: CANADA, British Columbia, on a southwest facing, open, wet hillside in Sooke Potholes Provincial Park beside the Sooke River, elev. 75 m, 48°24’N 123°43’ W, 1 May 1991, Benedict 28 V207976 (holotype: UBC). Herba annua obligata, a Mimulus guttatus DC. Pistillo 5-13 mm longo, corolla 6—20 mm longa et pistillo calycem aequante vel paulo longiore 2012] FIG. 2. differ; a foliis non bullatu, et caulus non alatis differ; planta tetraploidea. Annual or winter annual herb, bearing oppo- site pedicillate basal leaves graduating into sessile cauline leaves, 5—25 cm high, glabrous to minutely pubescent. Roots fibrous. Leaves with leaf blade palmately veined, regularly denticulate, widely ovate, apex obtuse to acute, 0.5—3 * 0.5— 2.5 cm becoming gradually reduced up the stem; leaf blade above adaxially green, frequently with anthocyanic spotting, glabrous to minutely pu- bescent, veins often purplish red near leafbase; leaf surface below abaxially silver-green to purple, glabrous, veins green. Petiole 0—2 cm BENEDICT ET AL.: SHY MONKEYFLOWER-A NEW POLYPLOID MIMULUS ei) ose ae oe F wy . se as " te 4, % ‘ : ms. ; 7 < * » i : Pics . ee eee A. ee , sage . - 4 Meiotic chromosome counts in Mimulus. A. M. sookensis (LSN), with 28 bivalents as seen in prophase I of meiosis. B. M. sookensis (NHI), shown with two daughter cells at late telophase I. Upper cell has 28 distinguishable univalents, while the lower cell has ca. 28 univalents. C. M. nasutus (TRT), with 14 bivalents at prophase I of meiosis. D. M. sookensis (ROG) as seen at late telophase I of meiosis,with two daughter cells each possessing 28 univalents. long, green-white to red-white; glabrous. Stems tending to quadrangular but not winged, <2 mm wide. Inflorescence few flowered to racemose, terminal, with 1 primary raceme, occasional secondary racemes arising from leaf axils, flowers opposite in leaf axils. Pedicel 3—22 mm long, red, glabrous. Calyx 5—13 mm long, central adaxial calyx lobe longer than other four, green, often with anthocyanic spotting, white hairs on margin, somewhat inflated upon maturity. Corolla bila- biate or sometimes cleistogamous, 5—22 x 2- 13 mm, yellow, corolla lobes subequal, palate densely hairy, red spotted, extending into tubes as two ridges, tube narrowly funnel shaped, 36 MADRONO 4-13 mm long. Stamens didynamous, upper stamens shorter, long stamens 4-12 mm. Pistil 5—13 mm; style white, minutely pubescent; stigma yellow, usually slightly exserted from calyx; ovary 2-5 mm, green; stipe 0-1 mm; stigma lobes may be thigmotropic. Capsule dehiscing by longitudi- nal slits with persistent style, crowned by a persistent calyx; lower calyx lobes curved up- wards toward upper calyx lobe upon maturity. Seeds up to 300 per capsule, oval, brown, 0.5 X 0.2 mm. Chromosome number tetraploid, 1 = 28. Found on wet, sunny, hillsides, cutbanks, and ditches on Vancouver Island and the Gulf Islands, British Columbia, on the San Juan Islands of Washington state, in the Willamette and Umqua River Valleys in Oregon, and also in one known site in Dos Rios, Mendocino Co., California, from sea level to 600 m. Flowers from late March to May. The species is named after Sooke Potholes Provincial Park on Vancouver Island where it was found to grow abundantly and where the type specimen was collected. The common name shy monkeyflower is suggested, because this monkeyflower disguises itself as M. nasutus, and the flowers are small, in contrast to the ‘gay’ and gregarious flowers of M. guttatus (Vickery 1952). Additional M. sookensis Specimens Examined CANADA. B.C.: Lasqueti Island, Trematon Mountain, 19 May 1985, Ceska 19167 (V 144698); N. Pender Island, Oak Bluffs, 4 Apr 1983, Ceska and Olgilve 14245 (V_ 133335); Saltspring Island, 5 1/2 km SW of Ganges, Lot 34, 18 April 1976, Douglas 9716 (V_ 136977): Saltspring Island, clearing at the end of Isabella Road,18 May 1980, Benedict 3 (UBC 207936); Mayne Island, Heck Hill, open bluff, 13 March 1980, Janszen 1532 (V 107521) and 6 Apr 1979, Janszen 978 (V 98035); Galiano Island, 12 May 1975, Wood 13 (V 97333); Galiano Island, west- facing slope overlooking ocean, Bluffs Park, 19 May 1993, Benedict 35 (UBC 207931); Gabriola Island, 21 May 1951, Raymer 5603135 (UBC 70999); Vancouver Island, Gonzales Hill near Victoria, April 1916, Newcombe s.n. (V 42590); Vancouver Island, Alberta Head, Newcombe s.n. (V 42592); Denman Island, wet cliffs facing Hornby Island, 7 Jul 1952, Brink s.n. (UBC 68843); Vancouver Island, Durrance Lake drain- age on rock outcrop, 9 May 1963, Young 63 (UBC 108599); Vancouver Island, Ucluelet, rocky ledges, 23 May 1975, Rose 75-284 (UBC 177970); Vancouver Island, Anderson Hill in Victoria, 17 May 1950, Krajina and Spilsbury s.n. (UBC 55012); Vancouver Island, Mount Wells, 8 mi W of Victoria on moist rocky cliffs, 12 May 1975, Calder and Taylor 20776 (UBC 80960); Vancouver Island, Esquimalt, 17 Apr 1917, Darling s.n. (UBC 45840); Vancouver Island, [Vol. 59 Victoria, 4 March 1912, Henry s.n. (UBC 80455); W slope of Mount Maxwell, Saltspring Island, 15 May 1963, Young 159 (UBC 221634); Vancouver Island, 5 km N of Cowichan Lake, 19 May 1990, Benedict 4 (UBC 207937); Vancouver Island, Nanoose Hill, N of Nanaimo, | May 1990, Benedict 1 (UBC 207934); Vancouver Island, Finlayson Arm Road, near Goldstream Provin- cial Park, 17 May 1990, Benedict 2 (UBC 207910); Vancouver Island, south slope of Observatory Hill, Saanich Peninsula,l May 1991, Benedict 27 (UBC 207935). USA. ORE- GON. Josephine Co.: above Rogue River 0.7 km W of entrance to Indian Mary Park, 3 May 1993, Strayley 7506 (UBC 208478); N of Grant’s Pass near South Hill summit, 13 Apr 1991, Benedict 23 (UBC 208138). Lane Co.: S facing road cut on N side of Dorena Lake, 6 Apr 1991, Benedict 11 (UBC 207932); Douglas Co.: Umpqua River Valley, 6 Apr 1991, Benedict 26 (UBC 207995); Umpqua Valley, Roseburg Quadrangle, July 1914 Cusick 4178a, (UBC 149306); Umpqua River, 21 mi below Umpqua, 20 May 1954, Steward 6641, (UBC 197132). WASHINGTON. San Juan Co.: rock outcropping on Orcas Isl., 13 Apr 1975, Gates 4, (UBC 263239). Gabriola Island, 21 May 1951, Raymer s.n. (UBC 5603135). Features Distinguishing M. sookensis and M. nasutus Mimulus sookensis is exceedingly similar in floral morphology to M. nasutus (Fig. 1). All characters overlap to a degree with M. nasutus, but under favorable growth conditions, the following structures tend to be more reduced in M. sookensis (M. nasutus measurements are presented here in parentheses): stem width <1] mm (<4 mm), calyx length 5-13 mm (6—16.5 mm), leaves 0.5-3 X 0.5-2.5 mm (0.5-10 X 0.5-— 7.5 mm), height 3—25 cm (5-50 cm), pedicel length 3—22 mm (4-26 mm), stipe length 0-1 mm (0.52 mm). Mimulus sookensis tends to have a longer pistil relative to its calyx and the difference in calyx and pistil lengths range from 2.5—3.5 mm (O—6 mm). The ratio of the width of the flower to the base in M. nasutus is usually >2 (<2). Mimulus nasutus often tends to have a more sharply angled and winged stem and the leaves are often bullate, while M. sookensis tends to have anthocyanic red spotting on the calyx more frequently than M. nasutus. Relationships and Distribution The genus Mimulus contains well over 100 species of monkeyflowers, and within the Simiolus clade, there are approximately 16—24 species, in- cluding M. guttatus, M. nasutus, and M. sookensis (Grant 1924; Pennell 1951). Comparable to the 2012] SON 48N 46N 44N 42N 40N 129W 127W 125W Fic. 3. locations in which M. sookensis has been recorded. rest of the genus, M. guttatus and its close allies are an exceedingly phenotypically and ecological- ly diverse group, making the M. guttatus complex and its close relatives an attractive system for ecological and evolutionary studies (Wu et al. 2008). Consequently, defining species relation- ships in this group of closely related monkey- flowers is challenging. As defined by Vickery (1978), the M. guttatus species complex is comprised of the common yellow monkeyflower, M. guttatus, and its close relatives, M. nasutus, M. laciniatus A. Gray, M. platycalyx Pennell, and M. glaucescens Greene. Pennell (1951) included a number of other taxa in the complex, including M. nudatus Curran, a linear-leaved serpentine endemic, and M. pardalis Pennell, a distinct form of monkeyflower with a prominently purple- spotted calyx, thought to be closely related to M. nasutus (Pennell 1947). A copper mine endemic, M. cupriphilus McNair, was later included in the complex (McNair 1989). Wu et al. (2008) recognize M. guttatus, M. nasutus, M. 123W British Columbia, CAN Washington, USA California 121W 119W 117W Geographic distribution of M. sookensis in western North America, with filled squares indicating laciniatus, M. platycalyx, M. glaucescens, M. cupriphilus, and M. nudatus as members of the M. guttatus complex at the rank of species. We suggest the addition of M. sookensis to this species complex. Based on present observations, it appears that M. sookensis is characterized by a disjunct distribution. In the northern portion of its range, M. sookensis is found throughout the southern end of Vancouver Island, British Columbia, in the Gulf Islands of British Columbia, including but not limited to Saltspring, Mayne, Galiano, Denman, Lasqueti, and Pender Island, and also on the San Juan Islands of Washington (Fig. 3). In the southern portion of its range, M. sookensis is found in the Willamette and Umpqua River Valleys of Oregon, and also in northern Califor- nia. In Oregon and California, collections are known from as far north as Mehama, in Marion Co., Oregon, and as far south as Dos Rios, in Mendocino Co., California (Fig. 3). It is con- ceivable that many more undiscovered M. 38 SON 48N — el A6N ae ews 44N 42N 40N 38N 36N 34N MADRONO . 128W 126W 124W 122W 120W 118W [Vol. 59 ali it 116W FiG. 4. Approximate location of M. nasutus, M. guttatus, and M. sookensis throughout western Washington, western Oregon, and California. U.S. counties where M. sookensis but not M. nasutus has been observed are filled in black, counties where M. nasutus but not M. sookensis has been observed are filled in grey, while counties where both species have been observed have diagonal hatching. Counties where M. guttatus has been observed in Washington are indicated with vertical hatching. sookensis localities exist throughout the northern and southern portion of its range. To illustrate the extent of field observations, which suggest an absence or rarity of M. sookensis throughout much of California, we have recorded the locations of M. nasutus collected in California (Fig. 4) that were used in either crossing, genetic, or flow cytometry analyses (see Table 3 and references therein). If WM. sookensis existed further south of Dos Rios, it is likely that it would have been mistakenly collected as M. nasutus, and subsequent analyses would have revealed its tetraploid nature. In mainland Washington state, no M. sookensis have been observed to date. Kiang and Hamrick (1978) were unable to find any M. nasutus in the Cascades of northern California, Oregon, and Washington. Additional evidence, based on recent collections in Washing- ton state, suggests M. nasutus is rare in Washing- ton, unlike M. guttatus (D. Lowry, Univ. of Texas-Austin, and C. Wu, Univ. of Richmond, personal communication). At many M. guttatus sites in Washington, neither M. nasutus nor M. sookensis has been observed (Fig. 4). This pattern suggests that both M. nasutus and M. sookensis may be rare in Washington state, or at the very least, that M. guttatus and M. nasutus do not commonly co-occur in this region, to our knowl- edge. If the rarity of co-occurrence of the two progenitor taxa in Washington state is a real phenomenon and not an artifact of sampling, the limited opportunities for hybridization between M. guttatus and M. nasutus in this region may in part explain the fact that M. sookensis is even more rare than M. nasutus in this region, and perhaps does not occur at all. We cannot exclude the possibility that isolated or ephemeral allotetraploids derived from M™. guttatus and M. nasutus are found elsewhere where M. guttatus and M. nasutus co-occur and may potentially hybridize. However, determining the exact range limits of M. sookensis is beyond the scope of this paper, and we present here simply what is known at this time regarding the 2012] BENEDICT ET AL.: SHY MONKEYFLOWER-A NEW POLYPLOID MIMULUS 39 TABLE 3. LIST OF LOCALES USED TO ILLUSTRATE LOCATIONS OF M. NASUTUS, M. SOOKENSIS, AND M. GUTTATUS, THAT HAVE BEEN CONFIRMED TO BE OF DIPLOID OR TETRAPLOID NATURE, THROUGH EITHER GENETIC ANALYSES, CROSSING EXPERIMENTS, CHROMOSOME COUNTS, OR FLOW CYTOMETRY. The locale ID may refer to: 1) the culture number given in a published chromosome count, 2) an examined herbarium specimen accession number or collector number, or 3) the ID given to the locale when published. Abbreviations used: na = not applicable. Species Locale ID County, State Reference M. nasutus 16 Calaveras Co., CA Benedict 1993 KIN Fresno Co., CA Sweigart and Willis 2003; Sweigart et al. 2007 SNF Fresno Co., CA Sweigart and Willis 2003 na Fresno Co., CA Kiang and Hamrick 1978 KNR Humboldt Co., CA Modliszewski and Willis (unpublished data) Cult. No. 6060 Inyo Co., CA Mia et al. 1964 na Kern Co., CA Kiang and Hamrick 1978 na Sierra Co., CA Kiang and Hamrick 1978 NMD Solano Co., CA Sweigart and Willis 2003 CMF Sonoma Co., CA Modliszewski and Willis (unpublished data) KRR Sonoma Co., CA Modliszewski and Willis (unpublished data) Cult. No. 5865 Sonoma Co., CA McArthur et al. 1972 M12 Tehama Co., CA Sweigart and Willis 2003; Sweigart et al. 2007 TOK Tulare Co., CA Sweigart and Willis 2003 na Tulare Co., CA Kiang and Hamrick 1978 Cult. No. 5327 Tuolumne Co., CA Mukherjee and Vickery 1962; n = 13 NDP Tuolumne Co., CA Sweigart and Willis 2003; Martin and Willis 2010 MEN Tuolumne Co., CA Sweigart and Willis 2003; Martin and Willis 2010 NCL Tuolumne Co., CA Sweigart and Willis 2003; Sweigart et al. 2007 NFN Clackamas Co., OR Modliszewski and Willis (unpublished data) HCN Josephine Co., OR Modliszewski and Willis (unpublished data) TRT Multnomah Co., OR _ See text SF Wasco Co., OR Fishman and Willis 2001; Sweigart and Willis 2003; Martin and Willis 2010 WSK Klickitat Co., WA Modliszewski and Willis (unpublished data) CLR Klickitat Co., WA Sweigart and Willis 2003; Sweigart et al. 2007 M. sookensis BVN Douglas Co., OR Fig. 3 WBP Douglas Co., OR Fig. 3 Benedict 207995 Douglas Co., OR See text ROG Josephine Co., OR See text; Sweigart et al. 2008 Strayley 208478 Josephine Co., OR See text Benedict 208138 Josephine Co., OR See text DRN Lane Co., OR Sweigart et al. 2008 HIL Lane Co., OR Fig. 3 LSN Lane Co., OR See text; Sweigart et al. 2008 PSG Lane Co., OR Sweigart and Willis 2003; Sweigart et al. 2008 SPB Lane Co., OR Sweigart et al. 2008 Benedict 207932 Lane Co., OR See text SAN Marion Co., OR Fig. 3 WTU 263239 San Juan Co., WA See text NDR Mendocino Co., CA Sweigart and Willis 2003; Sweigart et al. 2008 M. guttatus WSKG Klickitat Co., WA Modliszewski and Willis (unpublished data) RFA Lewis Co., WA Modliszewski and Willis (unpublished data) HAM Mason Co., WA C. Wu, personal communication HOC Mason Co., WA D. Lowry, personal communication CHR Pierce Co., WA C. Wu, personal communication AWP Skagit Co., WA Modliszewski and Willis (unpublished data) NCG Whatcom Co., WA Modliszewski and Willis (unpublished data) BRI Mariposa Co., CA Sweigart and Willis 2003; Sweigart et al. 2007 na Mariposa Co., CA Kiang and Hamrick 1978 NDR2 Mendocino Co., CA Sweigart and Willis 2003; Martin and Willis 2010 SHI Mendocino Co., CA Modliszewski and Willis (unpublished data) Cult. No. 5044 na MHA Cult. No. 5751 NBC Monterey Co., CA Plumas Co., CA Santa Clara Co., CA Santa Clara Co., CA Santa Cruz Co., CA Vickery 1955 Kiang and Hamrick 1978 Modliszewski and Willis (unpublished data) Vickery 1964 Sweigart and Willis 2003 AO MADRONO distribution of M. sookensis based on current collections. Interestingly, while M. sookensis commonly co- occurs with M. guttatus throughout its range, with few exceptions, in habitats where M. sookensis is present, M. nasutus tends to be absent. Mimulus nasutus and M. sookensis are known to co-occur at only two locations. Although Vancouver Island is at the northern limit of the range of M. nasutus, it is found to co-occur with M. sookensis at one site on the southern end of Vancouver Island (Nanoose Hill). This site is at a lower elevation than many of the other locations on Vancouver Island where only M. sookensis was observed (Fig. 3, Benedict 1993). The second site is along the Rogue River in southern Oregon; other M. nasutus sites have also been found in this region (Table 3). Additionally, at the southern periphery of the range of M. sookensis near Dos Rios, California, M. nasutus and M. sookensis are found within ca. 3 km of one another, but not within the same collection locale (Sweigart and Willis 2003). At present, there is insufficient evidence to determine whether or not the apparent absence of M. nasutus at many of the M. sookensis collection locales is a historical artifact or if the relative rarity of co-occurrence is caused by some unknown biological or abiological factor. DISCUSSION Within just the Simiolus clade of the genus Mimulus, there are over 21 well-documented occurences of polyploidy or aneuploidy (reviewed in Beardsley et al. 2004). The Mimulus glabratus heteroploid species complex in the Simiolus clade is characterized by ploidal races that are distrib- uted across a north-south latitudinal gradient (McArthur et al. 1972). Crossing barriers exist both between ploidal races, and to varying extents, within ploidal races (Alam and Vickery 1973; Vickery et al. 1976). Here, together with data from previous publica- tions (Fig. 2; Table 2; Sweigart et al. 2008), we have presented evidence of another instance of polyploid speciation—the previously undescribed M. sooken- sis. Although the triploid block is not absolutely complete between M. sookensis and its diploid progenitors, a triploid bridge is not likely to contribute significantly to gene flow or polyploid formation in a selfing taxa (Ramsey and Schemske 1998). Vickery found many other forms of polyploid and aneuploid monkeyflowers in the M. guttatus species complex during the course of his extensive cytogenetic work in Mimulus, but no record exists of M. sookensis (Mukherjee and Vickery 1959, 1960, 1962; Mia et al. 1964; Mia and Vickery 1968; Vickery et al. 1968; McArthur et al. 1972). Most of the autotetraploid M. guttatus that Vickery found were in the south- western U.S. (Arizona, Colorado, New Mexico, [Vol. 59 and Utah) and Mexico or in Alaska, but one autotetraploid M. guttatus was found in Multno- mah Falls, near Portland, Oregon. This individual was likely not M. sookensis, since Vickery’s identification indicates that it bore more resem- blance to M. guttatus than M. nasutus. Within the M. guttatus species complex, the autotetraploid M. guttatus subsp. haidensis Calder and Taylor is a distinct form of M. guttatus endemic to the Haida Gwaii (Queen Charlotte Islands) of British Columbia, Canada. Despite these autotetraploid forms of M. guttatus, M. sookensis will continue to remain a distinct species, due to the fact that the progeny of a cross between autotetraploids and allotetraploids will be tetraploid, and any back- crossing with a diploid will occur in the direction of the outcrossing species (M. guttatus), not the selfing species (M. sookensis). These backcross progeny, if existent, will likely be inviable or infertile, as was shown in Sweigart et al. (2008). Additionally, data from nuclear genes (Sweigart et al. 2008) does not show loss of M. nasutus gene copies, which would be expected if hybridization with autotetraploid M. guttatus had occurred. The newly described M. sookensis is broadly distributed in scattered locations throughout the valleys of western Oregon and northern Califor- nia, and also on the southern tip of Vancouver Island and the Gulf Islands of British Columbia and San Juan Islands of Washington. The seemingly disjunct distribution of M. sookensis raises the question as to whether or not the distribution is actually discontinuous, or if M. sookensis exists undiscovered in Washington; further field work in Washington could help to determine if the observed distribution is real. Data from plants of the Pacific Northwest suggest that the glaciations of the Pleistocene created discon- tinuous distributions that were later recolonized (Soltis et al. 1997). If MM. sookensis formed post- Pleistocene glaciation events, it may be that M. nasutus has yet to extensively recolonize Wash- ington state, in contrast to the more common ™. guttatus, and that the rarity of M. nasutus in Washington has contributed to more extreme rarity of M. sookensis in Washington. If ™. sookensis formed throughout the Pacific North- west prior to Pleistocene glaciations, it may have existed in glacial refugia on Vancouver Island and Oregon (Soltis et al. 1997; Brunsfield et al. 2001; Shafer et al. 2010), and has not yet extensively recolonized Washington. Of final note is the observation that Mimulus sookensis from different collection locations all appear to be phenotypically quite similar to M. nasutus. It would be interesting to know if M. sookensis was formed by multiple polyploidiza- tion events, as suggested by sequences from one of two nuclear genes sequenced to date (Sweigart et al. 2008), or if individuals from as far apart as British Columbia and California originated once, 2012] and then spread geographically to occupy their current distribution. If M. sookensis was indeed formed by multiple allopolyploidization events, as is common among polyploid plants (Soltis and Soltis 1993, 1999) it would be of great interest to know how these interspecific polyploid hybrids between M. guttatus and M. nasutus all came to have the appearance of M. nasutus. ACKNOWLEDGMENTS We thank James Beck for assistance with the chromosome squashes, Connie Robertson for help acquiring herbarium specimens, David Lowry and Carrie Wu for providing information regarding their field work in Washington state, Neil Jacobs for assistance with figure production, and members of the Willis lab (in particular, Elen Oneal, Lex Flagel, and Kathleen Ferris for insightful discussion of other Mimulus species). We also thank two anonymous reviewers for providing comments that strengthened the paper. This work was funded by a grant to Fred R. Ganders from the Natural Sciences and Engineering Research Council of Canada, an NSF Fibr awarded to John H. Willis, and a grant from the Oregon Native Plant Society awarded to Jennifer L. Modliszewsk1. LITERATURE CITED ALAM, M. T. AND R. K. VICKERY. 1973. Crossing relationships in the Mimulus glabratus heteroploid complex. American Midland Naturalist 90: 449-454. ANDERSON, L. E. 1954. 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In this authoritative but refreshingly slim volume, veteran botanist and plant ecologist John Sawyer describes the majestic landscapes and natural inhabitants of northwest California, an area that has occupied most of his professional career. Dr. Sawyer is an engaging and confident guide through the varied landscapes of northwest California, expertly weaving together physical and biological patterns with environmental and human history. As the author says in the opening pages, the book is not an encyclopedia of organisms of northwest California. Rather, it is a tour of an ancient and complex region that is at once at the edge of a state and a center of biodiversity in the western United States. The book is laid out in broad themes, first describing the geography of the region, next exploring major ecological themes in sequence, including patterns in vegeta- tion, environmental history, the evolution of species diversity, fire regimes and other agents of change, and the current and future conserva- tion status of the region. The book is an integration of a lifetime of ecological study and learning. One cannot study botany without becoming at least a little curious about geology, environmental history, and the effects of humans on the land, and Dr. Sawyer discusses all of these topics with uncommon ease and authority. Lovers of wildlife, on the other hand, may find this volume less rich than it could be, but will still find useful insights and information. Even the most casual visitor will notice that northwest California is diverse in its climates, landscapes, and vegetation types, so one must break up the landscape to discuss its natural history. The book takes the approach of dividing northwest California into two geologic regions, then into smaller landscape units called countries (e.g., the middle Sacramento country). The latter is unconventional, but appealing in its informal- ity and descriptiveness. This framework is used throughout the book to describe the various ecological phenomena, from geology to vegeta- tion to disturbance regimes. For lovers of botanical nuts and bolts, this is no regional manual, but the author does provide broad floristic information and species lists for selected habitats. He begins by presenting a counterintuitive but interesting and, I believe, accurate view of the regional vegetation patterns. Despite its well-known floristic diversity, he describes the region as being dominated by several climatic and elevationally driven zones that are dominated by just seven tree species. Upon this broad and deceptively simple canvas, however, subordinate species and hydrology, geology, and disturbance-associated microhabitats build im- pressive floristic detail. Moving from the general to the more specific, the book tells us of the major tree and shrub species in different ecological zones of the region, then adds greater detail along with descriptions of selected habitats, such as montane and subalpine meadows, serpentine and limestone outcrop areas, coastal environments, and wetland and riparian zones. Some descriptions are tanta- lizing. I personally cannot wait to see the ‘outrageous shrub diversity” on the Hosselkus Limestone of Shasta County. For decades, northwest California has been plagued by divisive perspectives on the effects of land use. However, Dr. Sawyer brings an unusually informed and moderate view to dis- cussions of forest management and the ecology of fire. Rather than providing hackneyed and ideological arguments for or against logging or fuels restoration, he often uses the findings from a surprisingly rich pool of primary studies in the region conducted by himself and his many student colleagues at Humboldt State University. (Graciously, he credits the students first when describing such collaborative projects.) The findings are very interesting and surprising. For instance, most Californians probably envision northwest California as a land of ancient and timeless forests, yet the author’s synthesis of paleoecology and disturbance ecology paints a story of a dynamic region that has rarely been in equilibrium, despite the venerable ages of some of its trees. Many seemingly ageless forest commu- nities have existed for just a few thousand years. Moreover, particular forest stands may owe their complexity not just to their age, but to a history of patchy fires. Still other forests may be surprisingly young and very different from the landscapes experienced and managed by the region’s native peoples just a couple of centuries ago. Visually, I found the book to be a bit wanting due to the decision to collect all the plates in the middle of the volume. The images themselves are lovely and descriptive, however. Also, the maps of each “country” in the opening chapters are exceedingly plain, consisting of a simple silver digital elevation model with a black polygon delineating the area of interest. Drawings or other illustrations are infrequent and unadorned. 2012] Nor does the writing in the book reach the artistic heights that readers have enjoyed in the writings of some California naturalists like John Muir, David Rains Wallace, or Elna Bakker. Nonethe- less, the prose is well-crafted and enjoyable. Despite these modest shortcomings, the book should be a valuable resource to anyone inter- ested in the ecology of northwest California. It summarizes over four decades of original re- search in a highly readable narrative, with sufficient tables and sources to serve as a useful reference. This slim, but substantive, book BOOK REVIEW 45 provides a great introduction to the general geography and ecology of the region and provides many interesting tidbits for those already living and working there. It would undoubtedly be a good backpacking companion to be savored by a campfire somewhere in the wilds of northwest California. The price, howev- er, at $75 for hardcover, might give pause to the zealous but cash poor. —DANIEL A. SARR, National Park Service, 1250 Siskiyou Ave., Ashland, OR 97520; dan_sarr@nps.gov. Volume 59, Number 1, pages 1-46, published August 17, 2012 SUBSCRIPTIONS — MEMBERSHIP The California Botanical Society has several membership types (individuals ($40 per year; family $45 per year; emeritus $32 per year; students $27 per year for a maximum of 7 years). Late fees may be assessed. 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