Madro鋘o; a West American Journal of Botany
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2 MADRONOVOLUME 59, NUMBER 3 JULY-SEPTEMBER 20 1 A WEST AMERICAN JOURNAL OF BOTANY PosTFiRE Chaparral Regeneration Under Mediterranean and Non-Mediterranean Climates Jon E. Keeley, C. J. Fotheringham, and Philip W. Rundel 109 Morphological and Isoenzyme Variation in Rhododendron OCCIDENTALE (WESTERN AzALEA) (SECTION PeNTANTHERA; Ericaceae) G. F. Hrusa 128 Measurement of Spatial Autocorrelation of Vegetation in Mountain Meadows of the Sierra Nevada, California and Western Nevada Dave A. Weixelman and Gregg M. Riegel 143 Reappearance of the Vanishing Wild Buckwheat: A Status Review OF Eriogonum evanidum (Polygonaceae) Naomi S. Fraga, Elizabeth Kempton, EeRoy Gross, and Duncan Bell ... 150 Ptychostomum pacificum (Bryaceae), a New Fen Species from California, Oregon, and Western Nevada, USA John R. Spence and James R. Shevock 156 A New Combination in Einanthus (Polemoniaceae) from Idaho AND Oregon y Joanna E. Schultz and Robert Patterson 1 63 Research & Recovery in Vernal Pool Landscapes C. Matt Guilliams 164 Montana 166 California 167 PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY Madrono (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbaria, Life Sciences Building, University of California, Berkeley, CA 94720. Subscription information on inside back cover. Established 1916. Periodicals postage paid at Berkeley, CA, and additional mailing offices. Return requested. Postmaster: Send address changes to Madrono, Kim Kersh, Membership Chair, Uni- versity and Jepson Herbarium, University of California, Berkeley, CA 94720-2465. [email protected]. Corresponding Editor—Matt Ritter Copy Editor—Richard Whitkus Biological Sciences Department Department of Biology Cal Poly, San Luis Obispo Sonoma State University 1 Grand Avenue 1801 E. Cotati Avenue San Luis Obispo, CA 93407 Rohnert Park, CA 94928-3609 [email protected] [email protected] Book Editor—Matt Ritter Noteworthy Collections Editor—Dieter Wilken Board of Editors Class of: 2012—Gretchen LeBuhn, San Francisco State University, CA Robert Patterson. San Francisco State University, CA 2013—Eric Roalson, Washington State University, WA Kristina Schierenbeck, California State University, Chico, CA 2014—Brandon Pratt, California State University, Bakersfield, CA Tom Wendt, University of Texas, Austin, TX CALIFORNIA BOTANICAL SOCIETY, INC. Officers for 2011-2012 President: V. Thomas Parker, Department of Biology, San Francisco State University, San Francisco, CA 94132, [email protected] First Vice President: Andrew Doran, University and Jepson Herbaria, University of California, Berkeley, CA 94720, [email protected] Second Vice President: Vacant. Recording Secretary: Michael Vasey, Department of Biology, San Francisco State University, San Francisco, CA 94132, [email protected] Corresponding Secretary: Anna Larsen, Jepson Herbarium, University of California, Berkeley, CA 94720, [email protected] Treasurer: Thomas Schweich, California Botanical Society, Jepson Herbarium, University of California, Berkeley, CA 94720, [email protected] 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, [email protected]; the Membership Chair, Kim Kersh, University and Jepson Herbaria, University of California, Berkeley, CA 94720, [email protected]; the Editor of Madroiio; and three elected Council Members: Chelsea Specht, Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720-2465, [email protected]; Ellen Simms, Department of Intergrative Biology, 1005 Valley Life Sciences Bldg., #3140, University of California, Berkeley, CA 94720, [email protected]. Staci Markos, University and Jepson Herbaria, University of California, Berkeley, CA 94720, [email protected]. Graduate Student Representatives: Genevieve Walden, Department of Integrative Biology and Jepson Herbarium, University of California, Berkeley, CA 94720, [email protected]. Administrator: Lynn Yamashita, University of California, Berkeley, CA 94720. [email protected]. Webmaster: Ekaphan (Bier) Kraichak, University of California, Berkeley, CA 94720, [email protected]. @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Madrono, Vol. 59, No. 3, pp. 109-127, 2012 POSTFIRE CHAPARRAL REGENERATION UNDER MEDITERRANEAN AND NON-MEDITERRANEAN CLIMATES Jon E. Keeley^ -, C. J. Fotheringham' - and Philip W. Rundel- 'U.S. Geological Survey, Western Ecological Research Center, Sequoia-Kings Canyon Field Station, Three Rivers, CA 93271 -Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095 [email protected] Abstract This study compares postfire regeneration and diversity patterns in fire-prone chaparral shrublands from mediterranean (California) and non-mediterranean-type climates (Arizona). Vegetation sampHng was conducted in tenth hectare plots with nested subplots for the first two years after fire. Floras in the two regions were compared with Jaccard's Index and importance of families and genera compared with dominance-diversity curves. Although there were 44 families in common between the two regions, the dominant families differed; Poaceae and Fabaceae in Arizona and Hydrophyllaceae and Rosaceae in California. Dominance diversity curves indicated in the first year a more equable distribution of families in Arizona than in California. Woody plants were much more dominant in the mediterranean climate and herbaceous plants more dominant in the bimodal rainfall climate. Species diversity was comparable in both regions at the lowest spatial scales but not at the tenth hectare scale. Due to the double growing season in the non-mediterranean region, the diversity for the first year comprised two different herbaceous floras in the fall and spring growing seasons. The Mediterranean climate in California, in contrast, had only a spring growing season and thus the total diversity for the first year was significantly greater in Arizona than in California for both annuals and herbaceous perennials. Chaparral in these two climate regimes share many dominant shrub species but the postfire communities are very different. Arizona chaparral has both a spring and fall growing season and these produce two very different postfire floras. When combined, the total annual diversity was substantially greater in Arizona chaparral. Key Words: Climate, dominance, fire, species diversity, spring and fall annuals. Chaparral is a fire-prone evergreen shrubland 1982) and occurs in widely disjunct patches from that is the dominant vegetation in the mediterra- near Prescott in the northeast to the southeastern nean-type climate (MTC) region of California mountains around Tucson and east to the (Keeley 2000). From a global perspective this southwestern edge of New Mexico (Cable 1975; vegetation is somewhat unique in that it not only Pase and Brown 1982; Whittaker and Niering tolerates frequent fires but many of the species have 1964, 1965). Arizona and California are broadly fire-dependent reproduction, similar to shrublands similar in the shrub dominants that are shared in other mediterranean-climate regions (Rundel between these two regions (Knipe et al. 1979). 1981; Keeley and Bond 1997; Keeley et al. 2005). Northeast Mexico chaparral is restricted to patches Chaparral shrublands, however, are not re- of severe substrate in the Sierra Oriental Moun- stricted to MTCs as this vegetation type is widely tains south of Monterey, and this vegetation shares distributed in southwestern USA and disjunct to some of the same shrub species and genera as the northeastern Mexico (Keeley and Keeley 1988). Arizona and California chaparral communities. Arizona mirrors the MTC in the winter rains that The Arizona and Mexican chaparral are of taper off to a late spring drought, but departs interest for what they can potentially tell us about from the MTC by addition of a second rainy the evolution of chaparral taxa. Paleoecological season in the summer. However, the importance studies have suggested that many chaparral shrub of summer rains to the dominant shrubs is a species originated in interior portions of the matter of some debate, as it appears that these southwest (Wolfe 1964, Axelrod 1989) and contem- rains play a minor role in shrub growth and porary populations in Arizona and northeastern reproduction (Vankat 1989). Northeastern Mex- Mexico are interpreted as remnants of a Tertiary ico has a winter drought and summer rain climate, chaparral like vegetation that comprises taxa that but the physiological responses of the shrub largely originated under non-medterranean type dominants to drought are remarkably similar to climates (Ackerly 2009; Keeley et al. 2012). CaHfornian shrubs (Bhaskar et al. 2007). Although postfire chaparral responses have Arizona chaparral covers more than two million been studied in great detail in the winter rain hectares (Schmutz and Whitham 1962; Bolander region of California, little is known about ' 110 MADRONO [Vol. 59 Table 1. Fifteen Dominant Plant Families in 600 I I Arizona and California Postfire Chaparral (a) P = 0.946 Sites Based on Aerial Coverage. 500 + State/family Normalized cover c o 400 Arizona -I—' V '' 03 Poaceae 1.00 Fabaceae 0.676 ;9- 300 o x" ">: Asteraceae 0.327 CD . .'. •. s_ Verbenaceae 0.138 200 Fagaceae 0.100 CO y y Molluginaceae 0.085 C13 Convolvulaceae 0.074 C 100 Liliaceae