VOLUME 58, NUMBER 4 OCTOBER-—DECEMBER 2011 EFFECTS OF ERADICATION AND RESTORATION TREATMENTS ON ITALIAN THISTLE (CARDUUS PYCNOCEPHALUS) Thomas McGinnis And Jon Keeley ....cccccccccsssccccccccccccccceeeeeeessescceseeeeeaaaanes 207 MORPHOLOGICAL ANALYSIS AND PHYTOGEOGRAPHY OF NATIVE CALAMAGROSTIS (POACEAE) FROM BRITISH COLUMBIA, CANADA AND ADJACENT REGIONS Kendrick L. Marr, Richard J. Hebda, and Elizabeth Anne Zamluk.......... 214 POPULATION ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER (PINUS BALFOURIANA) WITH A DISJUNCT DISTRIBUTION IN CALIFORNIA PQUrICIG TE, MGI OM Cy x, Peak ahotr tite ehec 8. nce taken Dass os ihe vie ROMY ese dS cin 234 MOLECULAR PHYLOGENETICS OF GARRYA (GARRYACEAE) BBW fat (a OLS 01/1 ge Tamer anentoee Sie A eRer MNES La NEREE. a Pmt NIOOT RE Ones OPT 249 LECTOTYPIFICATION OF ARCTOSTAPHYLOS HOOVERI (ERICACEAE) PDGVIG. SRC AAO ND sins Re beeriaCal ees toes ety etie Eee TSO RU 250 CALYPTRIDIUM PARRYI VAR. MARTIRENSE (MONTIACEAE), A NEW TAXON ENDEMIC TO THE SIERRA DE SAN PEDRO MARTIR, BAJA CALIFORNIA, MEXICO C. Matt Guilliams, Michael G. Simpson, and Jon P. Rebman................. 258 NOMENCLATURAL KANKEDORTS IN PHACELIA (BORAGINACEAE: HYDROPHYLLOIDEAE) Genevieve K. Walden and Robert Patt€rsOn ...........cccccccceeceuccneccnscesccusceecs 26) THE JEPSON MANUAL VASCULAR PLANTS OF CALIFORNIA, SECOND EDITION EN CHID CN eres Mert eialas ee hassthan nein oe ona sence base ea eerie see A cae eo ecanslies 213 OTROS ape =o Seen Ec Cee eR Ye ZI5 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. 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Alien forbs invade these grasslands, displacing both native and other alien species. Italian thistle is a noxious alien herb that has recently invaded these grasslands, including ungrazed blue oak (Quercus douglassii) and interior live oak (Quercus wislizenii) stands in Sequoia National Park. Here, Italian thistle tends to dominate under oaks and has the potential to substantially alter the foothill ecosystem by displacing native plants and acting as a ladder fuel that can carry fires into the oak canopy. We tested the effects of selectively reducing Italian thistle populations alone and in combination with restoration of native species. Two thistle eradication techniques (clipping and the application of clopyralid herbicide) and two restoration techniques (addition of native forb seeds or planting native grass plugs) were used. After two consecutive years of treatment we found: a) clipping was not effective at reducing Italian thistle populations (clipping reduced Italian thistle density in some areas, but not vegetative cover), b) herbicide reduced both Italian thistle density and vegetative cover for the first two growing seasons after application, but cover rebounded in the third growing season, c) native forb cover and species richness were not significantly affected by clipping or spot-treating with herbicide, d) the grass and forb addition treatments by themselves were not effective at reducing Italian thistle during the course of this study and e) sowing annual forb seeds after clipping resulted in greater forb cover and moderately reduced Italian thistle vegetative cover in the short term. Key Words: Alien thistle, annual grasslands, blue oak, foothills, Italian thistle, native plant restoration. Approximately 20% of California’s 6550 plant species are naturalized non-natives (http://ucjeps. berkeley.edu/interchange) and 200 of these are considered to be invasive, having the ability to displace native species and disrupt ecosystem processes (Bossard et al. 2000). Low elevation grasslands and savannas are among the most highly invaded ecosystems in the state, and today they are dominated by a few species of non-native Mediterranean grasses and a mixture of native and non-native forbs (Heady 1988; Bartolome et al. 2007). These grasslands are vulnerable to additional invasions by alien forbs. Some of the more noxious species are the spiny Asteraceae such as thistles in the tribe Cynareae (Bossard et al. 2006). Italian thistle (Carduus pycnocephalus L.), which is native to Europe and Asia, is a widespread noxious thistle with an annual or biennial life cycle in California annual grasslands and oak woodlands. It is particularly robust under blue oak canopies, suggesting it has high nitrogen and/or moisture requirements (Holm et al. 1997; Perakis and Kellogg 2007). Such invasions may threaten native understory species on these sites, which could be a _ particular concern if rare species occupy the same habitat. Dense Italian thistle populations with overlap- ping rosette leaves are capable of excluding native plant species by monopolizing light (Bossard and Lichti 2000). They also potentially affect wildlife movement due to deterrence from sharp spiny leaves and stems (Parsons 1973). Moreover, Italian thistle can potentially threaten oak trees, because, they can grow to 2 m and generate ladder fuels. Ladder fuels connecting surface litter to oak canopies during wildfires are a major contributor to blue oak canopy scorch and top- kill (Horney et al. 2002). The potential for Italian thistle to shade out native forbs, alter grazing patterns and convert surface fires into crown fires presents significant management concerns in the blue oak woodlands throughout the state. When caught during their early phase of colonization invasive plant eradication may be feasible, but once established, they present formidable challenges for resource managers (DiTomaso et al. 2007). Methods aimed at eradication of invasive species may produce short term reductions in cover and density, but populations typically return once direct control ceases. One reason is that eradication methods can disrupt ecosystem processes and create disturbance sites for future colonization of invasive species (D’Antonio and Meyerson 2002). Also, even with precise treatments, erad- ication methods affect potential native competi- tors as well as other invasive species (Di Tomaso et al. 2007). Non-native species thrive in Califor- nia annual grasslands partially because these 208 MADRONO © North Fork qoludy Sites oGoo — o =< c ie i] : x< = oy LL Ss © ) a Fic. 1. grasslands are highly disrupted ecosystems where the competitive balance between native species has diminished (Heady 1988). In the Sierra Nevada foothills of Sequoia National Park, Italian thistle has been a persis- tent invasive and the target of eradication attempts since 2002. Typically resource managers have utilized two eradication treatments: spray- ing with the herbicide clopyralid and cutting or pulling (http://www.nps.gov/seki/naturescience/ badcapy.htm). The present study was conducted to compare clopyralid and clipping and to couple ‘these treatments with two restoration treatments (sowing native forb seeds or planting native perennial grass plugs) meant to increase potential native competitors. We hypothesized that sup- pression of Italian thistle would be best achieved by coupling eradication with restoration of native competitors. Results were determined over mul- tiple years so that treatment longevity could be determined. METHODS Study Areas and Treatments Our study was conducted between May 2006 and June 2010 in the Kaweah River watershed in [Vol. 58 Italian thistle study sites, Sequoia NP, California. the Sierra Nevada foothills in Sequoia National Park (Tulare County, California; Fig. 1). The vegetation was blue oak and interior live oak woodland and savanna; here cool wet winters and warm dry summers characterize its Mediterra- nean climate. Snow is uncommon in the foothills of the southern Sierra Nevada and therefore most precipitation is from rain. Except for the winter of 2009-2010, precipitation was at or below the 57 year average during this study (Fig. 2). Using both National Park Service (NPS) maps and on-the-ground surveys of Italian thistles we established study sites throughout the 20 largest thistle populations in the park: 10 populations in the middle fork watershed and 10 populations in the north fork watershed. Within each of the two watersheds, 10 ‘“‘canopy”’ sites were selected beneath the drip-line of blue oak, interior live oak or California buckeye (Aesculus californica [Spach] Nutt.) trees, at least 1 m from a tree bole. An additional 10 sites were selected outside the drip-line and were considered ‘‘open”’ sites. The amount of shading and solar intensity in both canopy and open plots varied widely due to aspect, canopy height and proximity to trees, shrubs and boulders. Each population consisted of several thistle patches located both beneath trees and in the open. Because most Italian thistle 2011] 200 —— Study period ---- Average 1948-2005 ~€ 150 E c ry ie 2 ae a rs 100 r a / ! o ” ; / o 50 ; \ y / . ; ri 7 a ee 4 e eS SS SS YF & NE Re ERE | RS FIG. 2. S s& oy y S re + O McGINNIS AND KEELEY: MANIPULATION OF ITALIAN THISTLE 209 & a 100 = os. ad = 2 60 So bed =f 40 bed bed ae bed x 3 20 Hl fl ee « S g 9 S ¥ SO 2 ce & of & SS ES ES Rou e Se @ ae See ve ¢ Treatment under tree canopy B 70 ab om ac a cs i roy a acd ac S 5 30> BO 20'c| acd S @# ce Ns 10- de e a 0 | = , aa ce) S Q se) S x 50) S < SP & OS SF SOS S O & QS v % 3 Ss ss" © £ 2 O wo OS Treatment in open areas Fic. 3. 2009 Italian thistle density in canopy sites (A) >, p and open sites (B) with respect to combinations of Italian thistle eradication (clipping and herbicide treatments in 2007 and repeated in 2008) and site restoration techniques (seeding and planting once in the winter of 2007-2008) in Sequoia National Park foothills. The same letter above bars indicates no significant difference at « = 0.05. Native Grasses and Forbs Native grass plantings were not very successful due to extreme mortality in the first year, but native grass cover began to increase after the initial die-off. Only one third of plots that were planted had survivors after 1.5 years (June 2009), but one year later (June 2010) all of these plots still contained live native grasses. Native forbs occurred in all but one of the study plots before the treatments and in all plots after treatments. In canopy sites in 2009, native forb cover was not significantly different in seeded versus control plots. In open sites, native forb cover was not significantly affected by any of the treatments. Native forb cover was not assessed in 2010. Alien Grasses Alien grass cover was only significantly affect- ed by one of the treatments, and only in open McGINNIS AND KEELEY: MANIPULATION OF ITALIAN THISTLE Zl A D> 8 50 a ac 7 45 = _ 40 ac ad a . 35 ab ws 30 Do 25 bed 4 (] iw ee bd S { b bd So ce) S Q 9 S x 50) S < 2 sc Ss 2 roms & gh SF Wg Treatment under tree canopy B m fae < o le. 5 5 30 ab ab ap as ® 9 25 oe 7 os 1S s £10 =o be © » 5 5 Cc c a ° ° VV 50) S Q 5a) S A, se) S es XC Pr KM SK PM OL os a) rey , Ph ne ge ge o ©) xy Treatment in open areas Fic. 4. 2009 Italian thistle vegetative cover percentage in canopy sites (a) and open sites (b) with respect to combinations of Italian thistle eradication (clipping and herbicide treatments in 2007 and repeated in 2008) and site restoration techniques (seeding and planting once in the winter of 2007-2008) in Sequoia National Park foothills. The same letter above bars indicates no significant difference at « = 0.05. sites in 2009; clipping plus planting native grass significantly reduced alien grass cover in open sites in 2009. The five most abundant alien grass species (by cover) in these plots were Bromus diandrus Roth (most abundant), Avena barbata Pott ex Link and Bromus hordeaceus L. (both species were equally abundant and together covered 15% less ground surface area than B. diandrus), Avena fatua L. and Bromus arenarius Labil. (both equally abundant and comprised 40% less ground surface area than A. barbata and B. hordeaceus). Bromus diandrus was the domi- nant alien grass species in canopy sites and Bromus hordeaceous was the dominant alien grass species in open sites. Cover of the other abundant grass species was not significantly different between open and canopy sites. In general, clipping reduced alien grass cover while herbicide increased it. 212 MADRONO [Vol. 58 A 60 = : : gS 70> a 205 c | a MY ® 60 | a a a © 40 - -— | ab a ® ab | cc 2 & 40 4 ab ab § 30, b 6 2 30] , 7 8 10+ y © 10 - eo! } LU, | g a YY © & 8 £§ & & OH & 0 - mal: : = KE PK SF KF ST SK &” _ ) s oe ve ie gon ce Pre-treatment 2008 2010 g % vr S = ¢ SF Year Treatment under tree canopy FIG. 6. Italian thistle percentage vegetative cover in control (white bars), clipped (striped bars) and herbi- : Pf cide-treated (dotted) plots from 2007 (pre-treatment) s 60 through 2010. Canopy and open plots are combined for e P this analysis. The same letter above bars indicates no o SO 4 4 a a a significant difference at « = 0.05. rT) b= oo a B > 30 aes Daa = a Italian thistles in blue oak and interior live oak = 4 Ae a savanna plots in the short-term; however, it was «3 10 Fy not effective at controlling thistles two years after ° 0 ~ Nel treatments were stopped. Due to clopyralid’s pre- S y 9 S C&O emergent effect (DiTomaso et al. 2007) treat- 9 9 Treatment in open areas Fic. 5. 2010 Italian thistle vegetative cover percentage in canopy sites (A) and open sites (B) with respect to combinations of Italian thistle eradication Italian thistle eradication (clipping and herbicide treatments in 2007 and repeated in 2008) and site restoration techniques (seeding and planting once in the winter of 2007—2008) in Sequoia National Park foothills. The same letter above bars indicates no significant difference at = 0.05. Species Richness Native species richness the first year after all treatments were complete (2009) was not signif- icantly different between treatments and controls in canopy or open sites. The controls had a mean of 4-5 species per 4 m? plot, while the treatments had 4-7 species per plot. Treatments with the greatest mean native species increases (2 more native forb species per plot than the control) were: clipped plus seeded plots in canopy sites and seeded, clipped plus seeded, clipped plus planted and herbicide plus seeded plots in open sites. Compared to the controls in both the canopy and open, alien grass and forb species richness was not significantly affected by any of the treatments. DISCUSSION Spot-treating Italian thistle with the selective herbicide clopyralid was effective at controlling ments applied in the spring one year controlled Italian thistle density and cover the next year, yet the Carduus seed bank can remain viable for 8— 10 years (1.e., Burnside et al. 1996; Sindel 1997) and in order to be effective in the longer term, herbicide would need to be reapplied to draw down the seed bank. Unlike the herbicide treatment, clipping was not an effective eradica- tion treatment even in the short-term. Because Italian thistle seeds could eventually enter from outside sources and the fact that healthy native plant populations can resist aliens (Young et al. 2009), it has been proposed that eradication methods should be coupled with restoration treatments for the longest-lasting effect (D’ Antonio and Meyerson 2002; DiTomaso et al. 2007). We hypothesized that suppression of Italian thistle would be best achieved by coupling eradication with restoration of native competi- tors, but we did not find sufficient evidence to support this hypotheses. Although clipping fol- lowed by seeding native forbs modestly controlled Italian thistle density and cover in canopy sites in the short term, it did not have an effect two years after treatment. The other restoration treatment, planting native grasses, was also ineffective at reducing Italian thistle cover, at least in the few years of this study. Despite the low percentage of native grass cover and high mortality the first year, the fact that planting increased native grass fre- quency indicates that foothill sites currently devoid of native grasses could support these species, once established. Also, because peren- 2011] nial grasses grow so slowly in this environment longer term monitoring of this treatment might be useful. In these sites Italian thistle density and cover were stable over four years indicating that this species had saturated sites before our experiment began. Without intervention these populations are likely to remain as seed sources for further dispersal and will present formidable challenges for resource managers in the Sierra Nevada foothills. We recommend more research in the area of eradication combined with restoration so that an effective combination of treatments can be found. We learned that spot- treating with clopyralid was an effective short- term treatment, but we did not come up with a longer-term solution. Broadcast application of clopyralid would be easier to apply and would protect larger areas from reinvasion, but could potentially harm certain native plants; therefore, a study on the effects of this herbicide on native flora would be warranted. ACKNOWLEDGMENTS Thanks to Julie Yee (Statistician), Matt Brooks (Research Ecologist) and Todd Esque (Research Ecologist) from the USGS Western Ecological Re- search Center for their helpful comments and advice on this manuscript. We thank Melanie Baer-Keeley, Amy Brown and John Nelson for their many hours of hard work throughout the project. Athena Demetry and Anne Pfaff assisted in the planning and logistics. The following people helped with the field work: Corie Cann, Pam Collins, Linda Franson, Lori Gant, Lee Goldstein, Marcia Goldstein, Sarah Graber, Jonathan Humphrey, Audrey Lucterhand, Kelly McGinnis, Kate McLaren, Liz Nanney, Anne Pfaff, Shelley Quaid, Dylan Schwilk, Christine Shook and Rich Thiel. We also appreciate the following SEKI divisions that provided help: Interpretation, Resources Management & Science, and Maintenance. Funding was from USGS Park-Oriented Biological Support. Any use of trade, product, or firm names in this publication is for descriptive purposes only and does not imply endorse- ment by the U.S. Government. LITERATURE CITED BARTOLOME, J. W., W. J. BARRY, T. GRIGGS, AND P. HOPKINSON. 2007. Valley grassland. Pp. 371—397 in M. R. Stromberg, J. Corbin, and C. D’Antonio, (eds.), California grasslands: ecology and manage- ment. University of California Press, Berkeley, CA. BOSSARD, C., M. BROOKS, J DITOMASO, J. RANDALL, C. ROYE, J. SIiGG, A. STANTON, AND P. WARNER (eds.). 2006. California invasive plant inventory. California Invasive Plant Council, Berkeley, CA. AND R. LICHTI. 2000. Carduus pycnocephalus L. Pp. 86-90 in C. C. Bossard, J. M. Randall, and M. C. Hoshovsky, (eds.), Invasive plants of California’s wildlands. University of California Press, Los Angeles, CA. BURNSIDE, O. C., R. G. WILSON, S. WEISBERG, AND K. G. HUBBARD. 1996. Seed longevity of 41 weed species buried 17 years in eastern and Western Nebraska. Weed Science 44:74-86. McGINNIS AND KEELEY: MANIPULATION OF ITALIAN THISTLE 215 D’ANTONIO, C. AND L. A. MEYERSON. 2002. Exotic plant species as problems and solutions in ecolog- ical restoration: a synthesis. Restoration Ecology 10:703-—713. DITOMASO, J. M., S. F. ENLOE, AND M. J. PITCAIRN. 2007. Exotic plant management in California annual grasslands. Pp. 281-296 in M. R. Strom- berg, J. Corbin, and C. D’Antonio, (eds.), Califor- nia grasslands: ecology and management. Univer- sity of California Press, Berkeley, CA. 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TYSER, R. W., J. M. ASEBROOK, R. W. POTTER, AND L. L. KURTH. 1998. Roadside revegetation in Glacier National Park, USA: effects of herbicide and seedling treatments. Restoration Ecology 6:197—206. YOUNG, S. L.; J. N. BARNEY, G, B. KYSER, T..-5. JONES, AND J. M. DITOMASO. 2009. Functionally similar species confer greater resistance to invasion: implications for grassland restoration. Restoration Ecology 17:884—892. ZAR, J. H. 1990. Biostatistical analysis. Prentice-Hall, Englewood Cliffs, NJ. MADRONO, Vol. 58, No. 4, pp. 214-233, 2011 MORPHOLOGICAL ANALYSIS AND PHYTOGEOGRAPHY OF NATIVE CALAMAGROSTIS (POACEAE) FROM BRITISH COLUMBIA, CANADA AND ADJACENT REGIONS KENDRICK L. MARR Royal BC Museum, 675 Belleville Street, Victoria, B.C. Canada V8W 9W2 kmarr@royalbcmuseum.bc.ca RICHARD J. HEBDA Royal BC Museum, 675 Belleville Street, Victoria, B.C. Canada V8W 9W2 and Department of Biology and Schools of Earth and Ocean Sciences and Environmental Studies, University of Victoria, P.O. Box 1700, Victoria, B.C. Canada V8W 2Y2 ELIZABETH ANNE ZAMLUK' Western Edge Botany, Box 2085, Sidney, B.C. Canada V8L 3S3 ABSTRACT The taxonomically difficult and ecologically and phytogeographically important genus, Calama- grostis, was examined for British Columbia (BC). Morphological characters were analyzed by Principal Components Analysis (PCA) to characterize taxa and to aid in the development of a new key. Eight native species (Ca/amagrostis canadensis, C. lapponica, C. montanensis, C. nutkaensis, C. purpurascens, C. rubescens, C. sesquiflora, and C. stricta) are confirmed to occur in British Columbia, of which C. montanensis, C. nutkaensis, C. purpurascens, C. rubescens, and C. sesquiflora are reliably distinguishable. Comparison of species distribution to regional climatic and vegetation history suggests that Calamagrostis nutkaensis and C. sesquiflora likely survived in coastal refugia during late Wisconsin glaciations. Ca/amagrostis purpurascens likely persisted beyond the glacial limits or within nunataks and then spread into previously glaciated sites. Two interior continental species, C. montanensis and C. rubescens, probably spread north and west from the unglaciated zone south of the Cordilleran and Laurentide ice sheets. Calamagrostis lapponica likely persisted north of the ice sheets, and then spread southward into high-elevation sites in northern and eastern BC. Calamagrostis canadensis and C. stricta probably survived south and north of the ice sheets, and then spread into the previously glaciated terrain. Key Words: British Columbia, Ca/amagrostis, phytogeography, Poaceae, principal components analysis. Calamagrostis Adans. (reed grass) 1s a wide- spread northern hemisphere genus of approxi- mately 100 species (Marr et al. 2007) mainly of the temperate and Arctic zones (Hitchcock et al. 1969; Scoggan 1978; Tsvelev 1984; Greene 2001). Twenty-five native and one introduced species occur in North America north of Mexico (Marr et al. 2007). As circumscribed by Greene (2001), eight native species occur in British Columbia: Calamagrostis canadensis (Michx.) P. Beauv., C. lapponica (Wahlenb.) Hartm. C. montanensis (Scribn.) Scribn., C. nutkaensis (J. Presl) J. Presl ex Steud., C. purpurascens R. Br., C. rubescens Buckley, C. sesquiflora (Trin.) Tzvlev, and C. stricta (Timm.) Koeler. Two intraspecific taxa are recognized within C. canadensis and C. stricta. Species are primarily distinguished according to spikelet length, the length of the awn relative to the lemma, the position of attachment of the awn, Whether the awn is bent or straight, and callus 'Present address: 3040 North Road, Gabriola, B.C. Canada VOR 1X7. hair length relative to lemma length (Hulten 1968; Hitchcock et al. 1969; Clarke 1980; Tsevlev 1984; Marr et al. 2007). In 2005, Calamagrostis epigeios (L.) Roth, a Eurasian species, was collected for the first time from BC (British Columbia, Lower Mainland, Chilliwack, Fraser River, unnamed island in Fraser River N of Chilliwack, 49°12'18"N, 121°57'33”"W, 28 Aug 2005, Frank Lomer s.n. (V195593)). It and the recently introduced horticultural plant C. x acutiflora (Schrad.) DC., of Eurasian origin, are not included in this study (but see Marr et al. (2007)). Calamagrostis species in BC occur in diverse habitats including alpine tundra, coastal bluffs, wetlands, coniferous forest, steppe and meadows. Several species have prominent ecological roles because of their abundance, their characteristic associations with key regional ecosystems (Mei- dinger and Pojar 1991), and their ability to colonize following disturbance (Tsvelev 1984; MacDonald and Lieffers 1991). In the United States, natural stands of C. rubescens, C. 2011] montanensis, and C. inexpansa A. Gray (=C. stricta) provide forage, and C. canadensis 1s a source of wild hay (Hitchcock 1971). Specimen labels from DAO (Agriculture and Agri-Food Canada Herbarium, Ottawa, Ontario) indicate that in the 1970’s Agriculture Canada evaluated accessions of C. canadensis, C. purpurascens, and C. stricta from western Canada, in common garden plots in Beaverlodge, Alberta, presumably to bring these species into cultivation as forage. Calamagrostis sesquiflora is of notable phytogeo- graphical interest because its restricted distribu- tion may provide clues to the region’s glacial history and subsequent colonization by plants (Ogilvie 1997). For these reasons, the identification of Cala- magrostis species is particularly important, how- ever many have observed that species deter- minations are difficult. For example, Stebbins (1930:35) observed that Calamagrostis species ‘are exceedingly variable and difficult to define’, while Hitchcock et al. (1969:522) noted that ‘several species are highly variable and mutually distinguishable only with some difficulty’. In western Canada, the species of the ““C. canaden- sis/C. strictalC. lapponica complex” are especially challenging to distinguish. Within C. canadensis and C. stricta it 1s also difficult to assign many specimens to a subspecific rank sensu Greene (2001). In the Royal British Columbia Museum herbarium (V) many specimens were identified to the wrong species or subspecies, suggesting that published keys may not be adequate to separate taxa reliably. For example, many character states by which the subspecific taxa of C. canadensis and C. stricta are differentiated overlap greatly (Greene 2001). Classification of some North American species is difficult, in part, due to apomixis, hybridization and polyploidy (Nygren 1954; Clarke 1980; Greene 1984; Tsevlev 1984) which likely generate and perpetuate numerous closely related geno- types that differ from each other by relatively subtle differences. A description of the genus can be found in Marr et al. (2007). Multiple chromosome counts have been reported for the species that occur in BC (Nygren 1954; Kawano 1965; Moss 1983; Greene 1984): C. canadensis (2n = 42, 45, 48, 49, 51, 56, 62, 65); C. lapponica (2n = 28, 42, 49-140); C. montanensis ( 2n = 28); C. nutkaensis (2n = 28); C. purpurascens (2n = 40-42, 47-49, 50, 53, 56, 54, 84); C. rubescens (2n = 28, 42, 56); C. sesquiflora (2n = 28); C. stricta (2n = 28, 56, 70, 84, ca. 104, ca. 114, ca. 120, ca. 123). Polyploidy and especially aneuploidy occur in those taxa that are difficult to distinguish, namely C. canadensis, C. lapponica and C. stricta. We undertook a systematic examination of British Columbia Calamagrostis species, based largely on a multivariate analysis of morpholog- ical characters. Our goals were to develop a better MARR ET AL.: MORPHOLOGY OF CALAMAGROSTIS IN B.C., CANADA 215 key and species descriptions, to evaluate intra- specific taxa in C. canadensis and C. stricta, and to more accurately map species distributions. We examine the phytogeography of the genus in northwestern North America and discuss it in the context of current understandings of the regions glacial, environmental and climatic history (Hebda 1995, 1997; Byun et al. 1997; Whitlock and Bartlein 1997; Heusser et al. 1999; Heinrichs et al. 2002). The results presented here are the basis for the measurements and a portion of the key presented in the Flora of North America treatment for those Calamagrostis species that occur in British Columbia (Marr et al. 2007). METHODS Morphology We examined and recorded label data from 1900 specimens from multiple herbaria: V, DAO, CAN, OLYM, SMI, UAC and UBC. We concentrated on specimens from British Colum- bia, but included selected material from Alaska, Washington, Yukon, Alberta, and Russia (four specimens originally determined as C. langsdorfii (Link) Trin. that were annotated by C. W. Greene in 1991 as C. canadensis var. langsdorfii (Link) Inman). We also observed species habitats and sampled populations during fieldwork in British Columbia from 2002-2009. No type specimens were viewed. Under magnification, we measured and ob- served 24 characters (Table 1) from 247 speci- mens (Appendix 1), initially accepting as correct the most recent name on the sheet. Data were analyzed by Principal Components Analysis (PCA) using SYSTAT (Wilkinson 1997). Indi- viduals were plotted according to their scores from the first two PCA axes. Taxa whose specimens mostly grouped together and were distinct from other taxa in the scatter-plot were removed from the data set. The PCA was repeated a second time using the revised data set. By removing the specimens of the more distinct species we hoped to achieve some resolution among the specimens of the remaining taxa, 1.e., those that were less distinct in the scatter-plot. This procedure was repeated a third time. For the second and third PCA an additional character, “glume width’ (GW) was added because the ratio of glume length to glume width appeared to be useful to distinguish among the taxa included in these PCA’s. The character, ‘‘awn exserted versus not exserted” (AWNXRT), was removed for the second and third PCA’s because all specimens of the species included in these PCA’s shared the same character state (awns were not exserted). If a specimen’s position on the scatter-plot differed from others of the same taxon, or if it 216 TABLE 1. MADRONO [Vol. 58 CHARACTERS MEASURED ON SPECIMENS OF BC CALAMAGROSTIS TAXA FOR PRINCIPAL COMPONENTS ANALYSIS (MARKED BY “‘+”?) AND TAXONOMIC DESCRIPTIONS. Characters marked by ‘‘*’’ are those used by Greene (2001). Character Character code Character description Inflorescence +*INEL length +INFW width +RACH rachis surface: 1 = sparsely scabrous; 2 = very scabrous, with longer, twisted projections +*BRL longest branch from the most basal inflorescence node First glume +*GL length +GW width GLWRAT length/width ratio GVERSUSL glume length minus lemma length +GSR glume surface: 1 = glabrous; 2 = scabrous keels only; 3 = whole surface scabrous; 4 = scabrous projections longer and bent Lemma +LML length +HARL callus hair length *HRAT callus hair length/lemma length +AWNATT distance from base of lemma to point of attachment of awn AWBSRAT distance from base of lemma to point of attachment of awn/lemma length +* AWNXRT extent of awn exsertion beyond glume margin: | = not exserted; 2 = exserted +*DIR awn: | = straight; 2 = bent +AWNL awn length ALRAT awn length/lemma length Flower +ANTHL anther length Leaf +LFW width of second leaf below inflorescence +LFL length of second leaf below inflorescence LL length of longest leaf on specimen WL width of longest leaf on specimen +*BLADE leaf blade: 1 = involute; 2 = flat +LIGT ligule type: 1 = delicate and lacerate; 2 = stiff and not lacerate +*ULFS upper leaf surface: 1 = glabrous; 2 = slightly scabrous; 3 = very scabrous; 4 = scabrous + slightly pilose; 5 = tomentose +*LLFS lower leaf surface: 1 = glabrous; 2 = scabrous +*COLLAR collar: | = glabrous; 2 = scabrous; 3 = pilose; 4 = tomentose +LIGULE ligule surface: | = glabrous; 2 = short hairy; 3 = long hairy +LIGL ligule length Stem TOTAL HT total plant height +*HT height from root crown to base of inflorescence +CULM culm surface: | = glabrous; 2 = slightly scabrous; 3 = very scabrous +*NODE number of nodes (from the root crown to the inflorescence) had been collected outside of the main geo- graphical or ecological range of that taxon, it was examined more closely and often, but not always, re-determined as the taxon with which it clustered most closely. Using box plots of each character for each taxon, we noted those characters that overlapped relatively little among taxa and tested the possibility of using these characters as a means of distinguishing among taxa. We repeated these steps through several iterations to minimize the degree of overlap among clusters of the same taxon. This approach assisted us in the preparation of a key. The key was successfully verified in the field during 2002—2008. Once we had established the key, we examined all other specimens that had not been included in the multivariate analysis and made re-determinations as necessary. Mapping Latitude and longitude data from confirmed herbarium specimens were entered into a database. Where only place names were given, latitude and longitude were derived from maps, printed gazetteers (Canadian Permanent Com- mittee on Geographical Names 1985) and the web sites http://geonames.nrcan.gc.ca/ and http://geonames.usgs.gov/. After all specimens had been examined and annotated the database was updated with re-determinations. Records were then mapped using ArcView 9a (Environ- mental Systems Research Institute, Inc. 1992— 1999). The map projections used are an Albers Equal Area Conic (Sphere) with a central median of —115 degrees and reference latitude of 51 degrees. MARR ET AL.: MORPHOLOGY OF CALA MAGROSTIS IN B.C., CANADA 217 2011] 4 3 = eee 2 = “N oc or 2 E 1 : : Lis Le 0 -1 -2 -2 S S or wa O O — cr A At - © 1M Ag oO OA © i i FACTOR(1) FACTOR(1) Fic. 1. Principal Components Analysis of morphological characters of native British Columbia Calamagrostis species. Letters represents a single specimen: C = C. canadensis var. canadensis; A = C. canadensis var. langsdorfi; L = C lapponica, M = C. montanensis, N = C. nutkaensis, P = C. purpurascens, R = C. rubescens, Q = C. sesquiflora, 1 = C. stricta subsp. inexpansa, S = C. stricta subsp. stricta. The ‘‘*’’ indicates specimens that we have re-determined. Specimens labeled with ‘‘+”’ are from Russia. A. All species. B. Original data set but with C. montanensis, C. purpurascens; C. sesquiflora, and C. rubescens removed. C. C. canadensis, C. lapponica, C. stricta only. D: Same as Fig. 1C, but with specimens labeled according to the most recent name that was written on the label i.e., prior to this study. RESULTS among C. stricta, C. lapponica and C. nutkaensis. The first axis accounted for 24.8% of the varia- tion with inflorescence branch length (BRL), plant height (HT), and the number of nodes In the first PCA (all species), specimens of C| (NODE) contributing the most. The second axis montanensis, C. purpurascens, C. rubescens and to accounted for 19.2% of the variation with ligule a lesser extent C. sesquiflora formed largely type (LIGT), rachis surface roughness (RACH), discrete clusters (Fig. 1A). Calamagrostis cana- and glume surface (GSR) contributing the most. densis and C. nutkaensis also clustered separately The third axis (not shown) accounted for 14.5% to some extent. There was considerable overlap of the variation with glume length (GL), leaf Morphology 218 MADRONO [Vol. 58 TABLE 2. RANGE OF VARIATION IN MORPHOLOGICAL FEATURES OF CALAMAGROSTIS TAXA FROM BRITISH COLUMBIA. Unless otherwise noted, these characters were used in the Principal Components Analysis. See Table 1 for description of character codes. * = PCA2, PCA3 only; ~ = not included in PCA; # = PCAI only Calamagrostis taxon (n) AWNATT (mm) canadensis var. 0.3—1.0(1.6) canadensis var. Character canadensis (28) — langsdorfii (34) Inflorescence INFL (cm) (9)11-4(19) (8)9—15(24) INFW (cm) (1)2—3(7) (1.5)2.5-(8) BRL (mm) 29-45(57) (27)35—60(120) RACH (1)1.5(2) (1)1.5(2) First glume GL (mm) 2.5—.5(4.0) (3.5)4.0-4.5(5.2) ~GW (mm) (0.7)1.0-1.3(1.4) (0.7)1.0—1.3(1.6) *GLWRAT (2.1)2.6—3.2(4.0) (2.7)3.5-4.0(6.7) ~GVERSUSL — (0.0)0.3—0.6(1.6) (0.4)1.0—1.4(2.1) GSR 1—3(4) (1)2-4 Lemma LML (mm) (2.2)2.7-3.1(4.0) (2.3)2.5—3.0(4.0) ~HARL (mm) (1.7)2.5—2.9(3.1) (1.5)3.0—3.3(4.5) ~HRAT (0.7)0.9-1.1(1.4) (0.5)1.0—1.2(1.5) (0.3)0.5—1.3(1.7) ~AWBSRAT (0.1)0.2—0.4(0.7) (0.1)0.2—0.4(0.5) H#AWNXRT no no DIR 1 1 AWNL (mm) 1.92.6 (1,7)2=3.1 ~ALRAT (0.6)0.9-1.1(1.4) (0.8)1.0—1.1(1.4) Flower ANTHL (mm) — (0.8)1.2—1.3(2.0) (0.9)1.2—1.6(2.6) Leaf LFW(mm) (2)3—5(8) (2)3—6(10) ~WL(mm) 2.5—5(6) (2)4-7(11) LFL(cm) (11)16—24(41) (11)18—24(48) ~LL(mm) (9)22—31(40) (14)21—29(50) BLADE flat flat LIGT lacerate lacerate LIGL(mm) (1)4~-6(12) (3)5—8(12) LIGULE 1-3 (1)-3 ULFS (1)3-4 (1)3-4 LLFS 2 2 Stem HT (cm) (36)55—70(145) (18)50—90(154) ~TOTAL HT (50)65—80(160) (45)65—110(180) (cm) CULM 1—2(3) [? NODE (2)3-4(6) (2)3—5(7) lapponica (28) (4)8—1(16) (0.7)1-2(2.8) (21)25-35(54) 1(1.5) (3.6)4.0—5.0(5.4) (1.0)1.3—1.4(1.7) (2.6)3.0—3.5(4.4) 0.3—1.5(2.3(1.9)) (O)1-(2) (2.5)3.0—-3.8(4.7) (2)3.0—3.5(4.7) (0.6)0.8—1.0(1.2) (0.3)0.8—1.2(1.6) (0.1)0.2—0.4(0.6) no LQ) 1.4-3.1 (0.6)0.8—1.1 (1.1)1.3—1.7(2.0) (1.5)2.0—3.5(4.0) 2.5—3.5 (4)8—12(21) (10)13—17(26) flat usually entire (0.5)2—3(6) 1-(2) (1)2(4) | (8)20—40(69) (23)35—50(80) l 1—2(3) montanensis (20) (4)7-9(10) (0.7)1-2(2.5) 13-30 (1)1.5(2) (3.1)3.5-4.5(5.7) na na (0.3)0.8—1.2(2.1) 1-3(4) (2.7)2.9-3.5(3.8) (1.2)1.7—2.12.4) 0.4—0.8(1.3) 0.5—1.0(1.8) (0.1)0.2—0.3 sometimes 2 (1.0)2.0—3.1 (0.7)1.0—-(1.2) (1.1)1.8-2.4(2.5) (1)2+3) 2-3 (5)8-11(18) (10)12—19(23) usually folded entire (1)2-4(S.5) (1)2-3 (1)2-3 2 (9)20—25(44) 16—40(54) (1)2-3 ep nutkaensis (20) (8)13—3(31) (1.1)2-4.5(9) 27—70(105) 1(2) 4.4-6.0(6.6) (1.0)1.1-1.3(1.7) (3.4)4.0—5.0(5.5) (0.4)0.8—1.3(2.5) IK) (3.0)4.0-4.5(4.8) (1.1)2:0-2.5@.9) (0.2)0.5—0.7(1.0) (0.7)1.1-1.9(3.1) (0.1)0.3—0.4(0.5) no l 1.0—3.2 (0.6)0.7—0.9(1.3) (1.0)2.4—2.6(3.3) (2)4-10(13) (3)4-10(20) (4)18—40(52) (15)31-41(56) flat usually entire (0.5)2—3(5.5) 1-3 1-2 1 (31)45-—85(111) (42)55—105(135) 1-3 1—2(3) width (LFW) and anther length (ANTHL) contributing the most. In the second PCA, C. montanensis, C. purpurascens, C. rubescens and C. sesquiflora were removed from the data set. In this analysis, C. nutkaensis and to some extent C. lapponica were distinct. Calamagrostis canadensis and C. stricta were largely distinct at the species level; however, there was more overlap at the intraspe- cific level (Fig. 1B). The first axis accounted for 27.1% of the variation with number of nodes (NODE), LIGT and GSR contributing the most. The second axis accounted for 19.2% of the variation with GL, lemma length (LML) and ANTHL contributing the most. For the third PCA C. nutkaensis was removed from the data, leaving C. canadensis, C. stricta, and C. lapponica. Calamagrostis lapponica formed a relatively discrete cluster; however there was some overlap with specimens of C. stricta and C. canadensis (Fig. 1C). There was little overlap between C. canadensis and C. stricta. Relative to C. lapponica, and to a lesser extent to C. stricta, C. canadensis was the most variable species. The first axis accounted for 33.0% of the variation with NODE, LIGT, and LIGL contributing the most. 2011] purpurascens (21) (4)7-9(13) 0.9-2(2.8) 13—25(34) 2 — (4.5)5.5-6.5(7.4) na na (0.5)1.1—1.8(4.5) (1)2.5-4 (3.4)4.1-4.6(5.0) (0.9)1.2—1.5(2.4) 0.2—0.4(0.6) (0.3)0.5—1.0(1.4) 0.1—0.2(0.3) yes 2 (4.4)6.0—7.0(9.0) (1.0)1.5—1.8(2.2) (1.3)1.7—2.5(2.9) 2—3(5) (2)3—-5(6) (4)7—12(17) (11)22-27(42) flat or folded usually entire (3.5)2-4(9) 3 5 2 (20)35—55(70) (33)40—65(80) (1)2-3 (1)2(3) The second axis accounted for 15.3% of the sesquiflora (20) (4)7—-9(11) 0.8—2.5(2.8) 15—30 (1)1.5(2) (5.3)6.0—8.5(9.5) na na (0.5)1.3—2.5 |—2 (3.4)4.8—5.8(6.8) (0.8)1.2—1.8(3.0) 0.1—0.4 (0.5)1.0—1.5(2.5) (0.1)0.2—0.3(0.7) yes 2 (5.4)7.0—11.0(13.0) (1.3)1.6—2.1(3.5) (1.2)2.2—3.0(3.4) (2)3—5(6) (2)4-7 (3)8—12(18) (9)17—25(31) flat entire or lacerate (0.5)2—S(6) 1—2(3) 2 l 19—25(39) 30-40(46) 1(2) 1—2(3) TABLE 2. EXTENDED. Calamagrostis taxon (n) rubescens (18) (5)9—13(24) (0.7)1.5—(2.7) (12)20—0(100) I—1.5(2) (3.2)4.0-4.5(5.1) na na (0.5)1.1-1.9 1(2) 2.43.4 (0.7)1—1.4(2.3) 0.2—0.5(0.9) (0.3)0.4(1.2) 0.1—0.2(0.5) usually 2 2.1—3.0(4.4) (1.0)1.2—1.4(1.7) (1.0)1.3—2.0(2.6) (1)2-S(8) (1.5)3—5(8) (6)9-20(36) (12)24—6(42) usually flat lacerate (2)3—5(6) (1)2-3 1-3 12 (23)60—70(105) (50)70—90(126) 1(2) (1)2-3(4) stricta subsp. inexpansa (33) (6)8—11(18) (0.8)1—2(2.8) 16—50 1—1.5 3.0-4.0(4.8) (0.9)1.2—1.5(2.0) (1.9)2.5—3.0(3.6) 0.1—1.0(1.4) 1—2(3) (2.4)2.7—3.5(3.8) (1.8)2.3—2.9(4.2) (0.5)0.7—0.9 (0.2)0.5—1.2(2.3) (0.1)0.2—0.4(0.7) no 1(2) 0.9-2.6 (0.1).8—1.1(1.3) (0.9)1.5—1.8(2.4) (1.5)2—3(6) 2—5(6) (5)11—18(28) (9)15—24(34) usually flat usually entire (0.5)3-4(6) 1-3 (1)2-4 1-2 (22)35—65(88) (29)45—75(98) ]—2 1—3(4) MARR ET AL.: MORPHOLOGY OF CALA MAGROSTIS IN B.C., CANADA stricta subsp. stricta (17) (4)8—0(1 3) (0.7) 1—2(2.5) (14)20—25(33) 1-1.5 (2.2)2.5—3.0 (0.8)1.0—1.1 (2.0)2.4—-2.8(3.2) 0.1—1.5(1.6) 1—2(3) (1.9)2.2—2.5(3.0) (1.2)1.5—2(3.0) (0.5)0.7—0.8 (0.3)0.7—1.1(1.3) (0.1)0.3—0.5 no 1(2) 1.42.5 (0.7)1.0—1.2(1.5) (1.1)1.2—1.4(1.7) (1.0)2(2.5) 1.5-3 (9)18—25 (12)13—23(25) flat or folded entire (0.5)1—2(4) (1)2—3 (1)2-4 12 (27)35—60(94) (35)50—70(100) 1-2 1—3(4) ments were taken for the PCA as well as additional variation with GL, hair length (HARL) and LML contributing the most. In order to visually evaluate the impact of our analysis upon the identification of the “C. cana- densis/C. lapponicalC. stricta complex” we re- plotted the results of the third PCA, but labeled the points according to the most recent (i.e., prior to our analysis) identification on the sheet (Fig. 1D). The changes that are indicated in this figure include 29 redeterminations at the subspecific category and 32 redeterminations at the species level. Measurements and observations based on our species determinations, using only specimens from North America, are summarized in Table 2. These data derive from specimens for which measure- specimens that we observed in cases where those specimens that were measured for the PCA failed to capture values at the lower or upper end of the range of a particular character. In terms of overall stature (1.e., plant height, leaf size, and inflorescence size), C. nutkaensis and C. canadensis are the most robust of the B.C. species, and C. montanensis and C. sesquiflora are the smallest. Calamagrostis sesquiflora and C. purpurascens have the largest florets and the longest awns, and C. stricta and C. canadensis have the smallest florets and awns. Callus hairs are longest in relation to the lemmas in C. canadensis, C. stricta and C. lapponica and are shortest, usually less than half the length of the lemma, in C. montanensis, C. purpurascens, C. 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Of the second group, all but C. nutkaensis usually have exserted, bent awns. Specimens of five species appeared to be viviparous, 1.e., some florets contained “plantlets” rather than flowers. Those species and specimens are as follows: C. canadensis var. langsdorfii (Canada, British Columbia, Blue Canyon Creek, 15 Aug 1972, Taylor, Roy L., Beil, Charles E., Marchant, Christopher J., Oliver 6139 (DAO 199629)); Canada, Northwest Territories, Rein- deer Station, Inuvik, 31 Jul 1978, Fred Fodor 1336 (UBC 167838)); C. stricta subsp. stricta (Canada, Yukon, Burwash Landing, S. of Burwash Land- ing, Kluane L., 5 July 1944, Raup, Hugh M., Raup, L. G. 12261 (CAN 276421)); C. montanensis (Canada, Alberta, Cardstom, 2 Aug 1950, W.G. Dore 12258 (DAO 105913)); C. rubescens (Cana- da, British Columbia, Elko, Silver Spring Lake, S of third lake in Lakes chain, 21 July 1996, Roemer, Hans L. 96286A (V169178); Canada, Alberta, Waterton town site, ca. 3 miles NNE of, 9 July 1974, Douglas, George W., Douglas, Gloria G. 7954 (V069747)); Canada, British Columbia, Shuswap Lake, 19 July 1996, Martin, M.E. 1337 (V164711); Canada, British Columbia, Lone Butte, on Bridge Lake Road, 7 miles SE of 100 Mile House, 13 July 1956, Calder, J. A., Kukkonen, I., Taylor, R. L. 1S800 (DAO 106488); Canada, British Columbia 100 Mile House, 1 mi. N of 100 Mile House, 5 Sept 1954, Calder, J. A.; Savile, D.B.O.; Ferguson, J. M. 15485 (DAO 106489)); and C. purpurascens (Canada, Yukon, Carcross, 15 Aug 1960, Calder, J. A.; Kukkonen, I. 28289 (DAO 106364)). The label of V164711 bears the note “‘current year’s flowering spike pseudoviviparous previous apparently normal.” ao wetlands and prairie”’. lake bottoms to clear-cuts, burned over muskeg”’ and *“disturbed gravel pits. forests and ‘“Burned Disturbance 0-1800 Drained Elevation (m) 450-1478 Soils textured substrates several collections (sand to clay) or on gravel; notable association with limey and alkaline soils. peat; notable on Mainly clay to sand, alkaline sites. ' Strictly on fine Topographic setting wet bottomlands occasional on slopes. and flat terrain, terrain, rarely on Largely of damp, not Flat valley bottom slopes; mesic to mostly hygric. CONTINUED. TABLE 3. Associated plants Ecology and Distribution Populus balsamifera, and P. tremuloides. Carex notably associated, Salix spp. Pinus, Betula, Picea, Re-determinations of herbarium specimens, new collections and observations in the field helped refine the understanding of ecology (Table 3) and distribution of Calamagrostis species in the study region (Fig. 2). These results demonstrate that two taxa can occur virtually at the same site but in slightly different habitats and that the distributions of several taxa, in particular C. lapponica and C. sesquiflora, are more sharply constrained than maps heretofore (Douglas et al. 2002b:131—132) have shown. Most taxa favor open habitats such as meadows, grasslands, wetlands, tundra and shorelines in association with woody vegetation. Calamagrostis rubescens favors forest or parkland settings. Calamagrostis nutkaensis occurs in shaded forest settings, although it is mostly a species of openings. Considering moisture, there are two broad groups. Taxa of relatively dry sites include C. montanensis, C. purpurascens, C. rubescens, C. lapponica, and C. sesquiflora (but under a humid Habitat Commonly in meadows or grow at edge of rather than in wetlands; also in low thickets and generally open woods. rivers and streams; infrequent Moist meadows, fens, less on grassy slopes; noted to frequent in marsh and bog; occasionally with shrubs; grassland associated with full sun. Taxon stricta subsp. inexpansa stricta subsp. Calamagrostis stricta Calamagrostis 222 og JOO 200 i eee Ko lometers e 140 130 120 @ Calamagrosts fappasiica FIG. 2. MADRONO [Vol. 58 140 130 120 Distribution maps of species of Calamagrostis that occur in British Columbia. Maps do not indicate complete distribution of each species from areas adjacent to, but outside of British Columbia. For complete North American distributions see Marr et al. (2007). climatic regime). Taxa of relatively moist sites include C. canadensis (though not always in the case of var. canadensis), C. stricta and to a large extent C. nutkaensis. Two species, C. /apponica and C. purpurascens favor well to moderately drained crest and upper slope positions. Calamagrostis montanensis and C. rubescens occur mainly on well to moderately drained slopes or flat terrain. Calamagrostis canadensis occurs largely on lower slope and mesic to even hydric valley bottom sites. In the alpine of northern BC, we consistently find C. canadensis var. langsdorfii in moisture receiving sites, local depressions, or beside boulders or tree islands that trap winter snow. Where C. /apponica grows near C. canadensis, C. lapponica always occurs on a higher slope position and in drier sites. Calamagrostis mon- tanensis can grow near C. canadensis at the northern limits of its range, but C. montanensis occupies warmer, more open and drier sites than C. canadensis. Calamagrostis stricta favors mesic to hygric base-of-slope and valley bottom sites. Calamagrostis nutkaensis exhibits the widest range of conditions, from relatively dry ridge crests to valley bottom hygric moisture regimes, but is found only in the generally moist coastal climate. The genus occurs over a wide range of substrate textures ranging from bedrock, to clay and peat. Two species, C. purpurascens and C. sesquiflora, favor bedrock or coarse textured substrates. Other species occasionally occur on these coarse substrates but are most abundant on medium to fine textures, especially silt to sand. Calamagrostis stricta and C. montanensis especially favor medi- um to fine textured soils. Generally the species grow over a wide range of soil chemistry, however, C. canadensis and C. sesquiflora are notably associated with acid sites, and C. stricta and C. purpurascens are associated with alkaline and limy soils. 2011] 140 130 120 OQ 100 200 400 a Kalometers 140 130 120 O 100 200 400 a ee Kilometers @ Calamagrostis rubescens Fic. 2. Continued. Species of Calamagrostis in western Canada, and in particular in British Columbia, have distinctive distributions (Fig. 2). Only two species, C. stricta and C. canadensis, occur throughout the 0 100 200 es Kilometers Fic. 2. Continued. MARR ET AL.: MORPHOLOGY OF CALAMAGROSTIS IN B.C., CANADA 225 140 130 120 yor) 20 EE ARalometers @ Calamagrostis sesquiflora province and range widely in northwest North America (see figures in Marr et al. 2007). Calamagrostis stricta ranges from Alaska and Yukon southward into the western United States 224 and eastward into Canada’s prairie provinces and adjacent U.S. states. In British Columbia, to the extent that the two subspecies can be reliably distinguished, C. stricta subsp. inexpansa has the wider range, occurring both in the interior as well as on the coast including the Queen Charlotte Islands. Calamagrostis stricta subsp. stricta ap- pears to be a strictly inland and relatively northern subspecies in British Columbia. The range dips southward in the Rocky Mountains, and it appears to be associated with cold continental climates. Globally C. stricta is a circumboreal/circumpolar taxon (Hultén 1968). Calamagrostis canadensis occurs throughout the study area from Yukon across British Columbia southward into the Pacific Northwest states of the U.S. The range extends eastward into the Northwest Territories, Alberta and across Canada and the northeast U.S. (Marr et al. 2007). Calamagrostis canadensis var. cana- densis is distributed throughout British Columbia except on the Queen Charlotte Islands and the extreme northwest. Calamagrostis canadensis var. langsdorfii 1s recorded from all parts of the province, including the Queen Charlotte Islands and the northwest. Two species, C. sesquiflora and C. nutkaensis, exhibit a strictly coastal distribution (Fig. 2): C. nutkaensis from California to coastal Siberia, C. sesquiflora from northern Vancouver Island to coastal northeast Asia (Hulten 1968; Hitchcock et al. 1969; Greene 1993). Calamagrostis nutkaen- sis occurs on or immediately adjacent to, the shoreline. Nearly all of the specimens from higher elevations were collected on the Brooks Peninsula (Ogilvie 1997) or on small islands near the mainland (e.g., Dundas Island). Plants from these locations are often shorter and have narrower leaves than those closer to sea level, however even at the higher elevations, there are specimens as tall as 1 m and with broad leaves. The distribution of C. sesquiflora is concentrated in the Queen Charlotte Islands, occurring elsewhere in BC only on the Brooks Peninsula, VI (Vancouver Island). This species was previously thought to occur as far south as Washington State, however plants from that area have recently been re-determined as C. tacomensis Marr and Hebda (Marr and Hebda 2006). Three of the four remaining species (C. lapponica, C. montanensis, and C. rubescens) do not occur on the coast (Fig. 2). The fourth, C. purpurascens is widespread and occurs almost exclusively east of the Coast Mountain and Cascade-Coast Mountain crest well into the continent (Fig. 2). In 2008 this species was collected from mountainous inland Vancouver Island for the first time. There is also a notable near-coastal site for the species at high elevation on the relatively dry, east side of the Olympic Peninsula in Washington State. MADRONO [Vol. 58 The distribution of C. rubescens is strictly southern and inland within the study area with the northernmost occurrence near Francois Lake, British Columbia at 54.1°N latitude. The range remains largely inland until southern California where populations occur along the coast (Greene 1993). Calamagrostis montanensis is largely a species of the continental plains, common in southern Alberta and adjacent U.S. states east of the continental divide. In British Columbia it is considered rare (Douglas et al. 2002a), occurring in the northernmost extension of the plains along the Peace River near Fort St. John and in the southern Rocky Mountain Trench where it has crossed the Rocky Mountain front. In British Columbia, C. lapponica, a largely circumpolar species, occurs mostly north of 55.2°N. There are, however, isolated and disjunct populations in west central Alberta between 51.8 and 53.2°N on high mountain tops. DISCUSSION Taxa of Northwestern North American Calamagrostis and Their Distinguishing Features Based on the multivariate analysis of morpho- logical characters two groups of species emerge: 1) those that are relatively distinct; and 2) those in which there is some overlap in morphological characters and that are therefore more difficult to identify. In the relatively distinct group are C. montanensis, C. purpurascens, C. rubescens, C. sesquiflora and C. nutkaensis. Those that are more difficult to correctly identify belong to the C. canadensis!C. lapponicalC. stricta ““complex”’. A comparison of Fig. 1C to Fig. 1D reveals that many specimens previously identified as C. stricta, C. canadensis or even C. purpurascens were re-determined in our analysis as C. lappo- nica, and vice versa. Further comparison of Figs. 1C and 1D indicates that the criteria that we used to distinguish between C. canadensis and C. stricta largely match the label determinations of the specimens at the species level, but less so at the infra-specific level. Although many specimens in this group are clearly distinct from each other, discrete clusters do not emerge in the scatter-plot of the PCA to the same extent as the other species. There are, however, morphological and ecological criteria by which these taxa can be distinguished and these are highlighted below. Calamagrostis lapponica vs. C. canadensis and C. stricta. We distinguish C. lapponica from C. stricta and C. canadensis largely according to habitat and lower leaf surface texture (Table 2). Both C. canadensis and C. stricta are species of | relatively moist even wet habitats (Crackles 1994; Cody 1996; Greene 2001), frequently occurring at | 2011] lower elevations than the largely alpine species C. lapponica (Table 3), although C. canadensis occa- sionally occurs in wet alpine meadows (and all of these are var. /angsdorfii). On one occasion we collected C. Japponica growing in standing water, a very unusual habitat for this species. The lower leaf surface of C. lapponica is glabrous, whereas in C. canadensis and C. stricta it is scabrous. The number of nodes is particularly useful to distin- guish C. /apponica, 1—2(3) nodes, from C. canadensis, (2)3—5(7) nodes, with which it is most frequently confused (compare Figs. 1C and 1D). Using the preceding criteria, the specimens that we considered to best fit C. lapponica grouped together in the PCA analysis (Fig. 1C) and the habitat of these specimens generally matches descriptions from other parts of the species’ range. In Europe, C. /apponica 1s largely an alpine species of “‘tundra, dry heaths and woods” (Clarke 1980). In Russia, C. /apponica occurs “‘In forest tundras, riverside sands and pebbles, sparse forests, among shrub; up to upper (bald) moun- tain peaks” Tsvelev (1984). Calamagrostis canadensis vs. Calamagrostis stricta. Criteria to distinguish C. canadensis from C. stricta are difficult to circumscribe because many characters overlap (Table 2, Fig. 1C). Using Fig. 1C as a starting point, we designated specimens plotted to the left of “O”’ on the first axis as C. stricta (or C. lapponica) and those to the right of the “0” as C. canadensis. The characters for which there was little overlap between species and which accounted for most of the variation in the PCA included NODE, LIGT, LIGL and BRL. Calamagrostis canadensis has (2)3—5(7) nodes, whereas C. stricta has 1—3(4) nodes. Ligules of C. canadensis are lacerate and fragile, whereas most C. stricta (and C. lapponica) specimens have relatively stiff ligules with entire margins. Calamagrostis canadensis ligules are generally longer ((1.5)3—8(12) mm) than those of C. stricta ((0.5)1—3.5(6) mm). There is little overlap in the length of the longest inflorescence branches (BRL), 1.e., less than 37 mm in 92% of C. stricta specimens and greater than 37 mm in 75% of C. canadensis specimens. Two ecological features also help to distinguish C. canadensis and C. stricta. In general, C. stricta appears to be associated with limey or alkaline substrates. In our experience, C. stricta is more likely to grow at the edge of, rather than in, wetland habitats as compared to C. canadensis. Where the two grow near each other C. stricta occurs “‘upslope” of C. canadensis and often at the edge or even just within forest, woodland or thicket. Recognizing subspecific taxa. Our analyses uphold the recognition of subspecific taxa in C. canadensis and C. stricta but reveal that the entities are part of a morphological continuum, a MARR ET AL.: MORPHOLOGY OF CALAMAGROSTIS IN B.C., CANADA 225 conclusion that matches that of Greene (1980). Most specimens of C. stricta in the lower part of Fig. 1C have glumes shorter than 3 mm long, whereas those that plot closer to C. /apponica, have glumes longer than 3 mm. This character primarily distinguishes subsp. inexpansa (glumes 3—4(4.8) mm) from subsp. stricta (glumes (2.2)2.5— 3.0 mm). Using this criterion C. stricta subsp. inexpansa occurs on the coast as well as inland, whereas C. stricta subsp. stricta is an inland taxon only (Fig. 2). There also appear to be habitat differences between the two, with subsp. inex- pansa growing in more forested habitats and subsp. stricta in more open habitats (Table 3). Hulten (1968) distinguished between these two taxa primarily by differences in callus hair length relative to the lemma, recognizing them at the specific rather than the subspecific level. He too mapped C. neglecta (Ehrh.) G. Gaertn. (C. stricta subsp. stricta) as primarily an inland species of coastal Alaska, and C. inexpansa(C. stricta subsp. inexpansa (A. Gray) C. W. Greene) along the coast as well as in the interior. Glume characters also help distinguish varie- ties of C. canadensis. Nearly all specimens of C. canadensis that plotted below *‘0” on the second axis (Fig. 1C) have relatively glabrous glumes, shorter than 3.5 mm. These we designated as C. canadensis var. canadensis (glume length (2.5)3— 3.5(4) mm). Those that occur above ‘‘0”’ have scabrous glumes longer than 3.5 mm, often with the scabrosity relatively long and sometimes bent. This second group we designated as C. canadensis var. langsdorfii (glume length (3.5)4—4.5(5.5) mm). All of the specimens labeled as C. langsdorfii from Russia are in this second group. The scabrosity of the glumes is mostly restricted to the keel in C. canadensis var. canadensis, whereas in C. canadensis var. langsdorfii they often cover the entire surface. Glume length to width differs as well, (2)2.5—3.5(4) for var. canadensis and (3)3.5—4(5.5) for var. langsdorfii, 1.e., glumes of var. /angsdorfii are more attenuate. The characters by which C. canadensis var. langsdorfii is recognized here are consistent with Hitchcock et al.’s (1969:525) view. They ques- tioned the occurrence of such an entity in North America, but concluded that if in fact it is the same as the Russian species, C. /angsdorfii, that it should then be included in C. canadensis var. scabra (Kunth) Hitche., which they distinguished primarily by the longer “rather strongly sca- brous” glumes. However, as Greene (1980) indicates, Calamagrostis scabra J. Presl was published in 1830, whereas C. /angsdorfii, was published in 1824. Our analysis (Fig. 1C) indi- cates that plants from Russia identified as “C. langsdorfiv’ are the same as C. canadensis var. langsdorfii. The use of the aforementioned characters to define intraspecific taxa for C. stricta and C. 226 canadensis generates better defined clusters in the PCA, though there remains overlap (compare Figs. 1C, 1D). Despite the overlap, we believe that continuing to recognize subspecific taxa is appro- priate, in part because the criteria that we used match subtle, but significant ecological differences (Table 3). Ecological differences are not so clear MADRONO [Vol. 58 in the case of C. canadensis subspecific taxa, nevertheless, there appears to be adequate mor- phological differentiation (glume size, shape and surface texture), to continue recognizing the two varieties. Whether or not these intraspecific taxa actually represent distinct lineages should be investigated using DNA markers. KEY TO CALAMAGROSTIS OF BRITISH COLUMBIA la. Awns more than 4 mm long (total length); glumes distinctly keeled; awns bent, exserted; upper leaf blade glabrous, slightly scabrous, or densely tomentose; inflorescence less than 13 cm long; 1-3 nodes......... Z 2a. Upper blade surface tomentose; leaves flat or involute; awns 4-9 mm long; mostly east of the Coast ee ee ee er ee oan ere eat C. purpurascens 2b. Upper blade surface glabrous to slightly scabrous; leaves flat; awns 5-13 mm long; Vancouver Island, Queen Charlotte Islands.................. C. sesquiflora lb. Awns less than 4 mm long; glumes keeled or rounded; awns straight or bent, if exserted, then less than 2 mm beyond lemma margin; upper blade glabrous to scabrous, never densely tomentose; inflorescence usually longer than 5 cm; 1-7 nodes. ........... Ce ae ee ar rae er nr re ere ee oe 3 3a. Callus hairs less than 60% of lemma length (observe here the callus, not the hairs of the rachilla) or the awn more than | mm longer than the lemma; glumes nearly glabrous; nodes 1—2(4). ............. 4 4a. Awns straight; plants strictly coastal, (within 10 km of coastline); collar not hairy; longest inflorescence branches 27—70 (105) mm long C. nutkaensis 4b. Awns bent; plants of the interior of the continent; collar often hairy; longest inflorescence branches (12)20—30(100) mm long. ....... Ses tit Gs Soa Adah as Aetie Se aan ee C. rubescens 3b. Callus hairs greater than 60% of the lemma length, if less than 60%, then the awn bent and less than 1 mm longer than the tip of the lemma or the glumes scabrous; nodes 1—5(7)................... 5 5a. Awns always bent, lower blade surface scabrous; glumes and lemmas scabrous; leaves (1)2(3) mm wide; inflorescence less than (4)7—9(10) cm long, the longest branches less than 30 mm long; 1—2 nodes; dry praimes-or grasslands, never alpime....2 os os aso dee es oe ee ee C. montanensis 5b. Awns nearly always straight; lower blade surface glabrous or scabrous; glumes and lemmas glabrous or scabrous; leaves 1-13 mm wide; inflorescence (4)8—-15(25) cm long, the longest branches 14-120 mm long; 1—7 nodes; forest, wetlands or alpine. .....................04. 6 6a. Longest inflorescence branches more than 37 mm long, or if shorter, then the ligule tip delicate (i.e., easily torn) and lacerate; glumes scabrous on the keel and often throughout, the projections sometimes bent; leaves flat, the lower surface scabrous; (2)3—5(7) nodes. ......... C. canadensis 7a. Glumes usually less than 3.5 mm long; glumes acute, and scabrous on the keels, rarely HhTouchout. «445.43 vac, eset sees Cae Se he Mel See eS Oe var. canadensis 7b. Glumes usually greater than 3.5 mm long, acuminate; glumes scabrous on the entire surface, the projections often bent. . ee ee we ee eee var. langsdor fii 6b. Longest inflorescence branches less than 37 mm long or if greater than 37 mm then the ligule stiff and entire; glumes glabrous to scabrous on the keels, but the projections never bent; leaves flat or involute, the lower surface glabrous or scabrous; 1—3(4) nodes. ............. 8 8a. Lower blade surface glabrous; leaves flat; glumes glabrous, rarely slightly scabrous on the keel, (3.5)4—5(5.5) mm long or more than 3 times longer than wide.......... C. lapponica 8b. Lower blade surface glabrous or scabrous; leaves flat or involute; glumes glabrous to scabrous, (2.2)2.5—4(4.8) mm long, usually less than 3 times longer than wide. ...C. stricta 9a. Glumes less than 3 mm long, glabrous to slightly scabrous and the callus hairs less than 2.5 mm long;-culms usually smooth: 0.22. <..0¢. 4-4 ss ee oe ee subsp. stricta 9b. Glumes more than 3 mm long, scabrous or the callus hairs more than 2.5 mm long; culms usually scabrous. ...... Distributions of Calamagrostis Species in BC and Their Phytogeographical Significance Extant Calamagrostis distributions can be understood in the context of Late Pleistocene history of habitats and glacial history. The most likely explanation for the strict coastal distribu- tion of C. nutkaensis and C. sesquiflora is that they survived on the coast during late Wisconsin (Vashon) glaciations both south and north of the Cordilleran Ice sheet or in shoreline refugia (closed refugia sensu Lindroth 1969). The habi- tats for both likely existed along the immediate re ee ee eee subsp. inexpansa shore zone even during the short interval of full- glacial conditions. Calamagrostis nutkaensis could have spread rapidly following deglaciation along the strandline and then moved a short distance inland into non-shoreline habitats such as bogs. Calamagrostis sesquiflora would seem to be well-adapted to persist on unglaciated head- lands and ridges such as envisaged in coastal refugia (Calder and Taylor 1968; Hebda 1997; Ogilvie 1997). However, its apparent need for | relatively mild winter temperatures and relative | drought intolerance (as evidenced by its oceanic distribution) prevented spread inland on what 2011] was a largely cold dry open landscape during glacial times (Hicock et al. 1982; Hebda and Whitlock 1997; Whitlock and Bartlein 1997; Heusser et al. 1999; Clague et al. 2004). The pattern of distribution of C. sesquiflora matches the disjunct ranges of other species with Vancou- ver Island/Queen Charlotte Island distributions such as Ligusticum calderi Mathias & Constance (Douglas et al. 2002a). Such distribution patterns are used as evidence of glacial refugia in these locations (Buckingham et al. 1995). The mostly interior, middle- to high-elevation C. purpurascens likely survived throughout the unglaciated, inland landscape and then spread into previously glaciated BC from circum-glacial refugia where suitable habitats for it occurred (Hicock et al. 1982; Ritchie 1987; Thompson et al. 1993). It may also have survived in high elevation nunatak refugia. A hardy species of mesic to xeric ridge tops and upper slopes, it would have been well suited to cold windblown full-glacial sites before 14,000 years ago. Though ice sheets covered much of British Columbia and Alberta, refugia occupied by non-arboreal vege- tation occurred, especially along the coast (Mathewes 1989; Hebda 1997; Brown and Hebda 2003; Fitton 2003) and possibly in northern B.C. (Marr et al. 2008). Both of the continental inland species, C. montanensis and C. rubescens, likely spread north and west from the unglaciated zone south of the ice sheets as suitable, relatively warm habitats became available. Considering the once greater extent of warm and dry climates and open habitats of the early Holocene (Hebda 1995; Heinrichs et al. 2002), their ranges, especially that of C. montanensis, was likely greater than today having shrunk with the forest expansion of the past 7000 years. The spread of C. rubescens has likely been limited by conifer forest development, but during the middle and early Holocene, when forests were much more open it likely grew further north and at higher elevations than it does today. The distribution and ecology of C. lapponica strongly suggest that it survived in the dry, cold, open landscape north of the Cordilleran and Laurentide ice-sheets during glacial times, and not south of these ice-sheets. Since the end of the ice age, it may have somewhat expanded its range southward, colonizing suitable high-elevation sites in the northern Rockies and further south into the Alberta Rockies. An alternative expla- nation for the Alberta sites would postulate that these populations are refugial relicts, remnants of a once more-continuous distribution. Extensive expansion after glaciation into southern alpine zones may not have been possible due to the rapid spread of conifers northward (MacDonald 1987). Our recent collections in northeast BC in the vicinity of Williston Lake and Tumbler Ridge MARR ET AL.: MORPHOLOGY OF CALAMAGROSTIS IN B.C., CANADA eae demonstrate that this species ranges further south in BC than was previously known. The wide-ranging C. canadensis likely survived south and north of glacial ice sheets and then reinvaded glaciated terrain. Calamagrostis cana- densis no doubt had abundant sites in which to thrive south of the Cordilleran-Laurentide ice system during the relatively moist Late Pleisto- cene time of extensive glacial lakes. This period of temporary separation may have contributed in part to the establishment or strengthening of differences between the two intraspecific taxa. Of the two subspecies, the hardier and widespread var. langsdorfii may well have survived in and spread from Beringia. With warming and moist- ening climates, C. canadensis, especially var. canadensis, may have been dispersed rapidly to every suitable wetland site by birds (adhering to feathers), thus spreading rapidly across the entire region after deglaciation. The wide-ranging distribution and similar, but not identical, habitats of C. stricta suggests a similar history to C. canadensis. However, the relatively-low upper-elevation distribution limit (Table 3) for C. stricta subsp. stricta suggests that it is less hardy than C. canadensis and may have come to occupy its North American range through northward migration from ice-free zones. The occurrence of C. stricta subsp. inexpansa as the only subspecific taxon on the coast is notable, because it suggests that the variety might be a pre- glacial entity isolated at one time from its vicariant partner, subsp. stricta. The distribution of subsp. stricta, coupled with the widespread occurrence until the Late Holocene of open habitats in the northwest interior of North America (Hebda 1995) strongly suggests a conti- nental interior source of spread for this taxon. CONCLUSIONS Our study of Calamagrostis in British Colum- bia upholds the taxonomic entities recognized previously for the province, but provides a more satisfactory treatment and key for this difficult grass genus in a phytogeographically critical region of North America. Our results demon- strate that combining morphologic, ecologic and distribution data can be an effective way of clarifying the taxonomy of a group of morpho- logically similar taxa. We suggest that major collections need to re-examined and annotated so that regional distribution maps can be corrected. For example, some of the previously published (Greene 2001) maps of Calamagrostis species distributions in BC were in error based on mis- identifications; in particular, the following chang- es should be made: C. /apponica does not occur near Smithers, BC or at the BC-Washington border; C. sesquiflora does not occur near Prince Rupert, BC; in southern BC, C. montanensis does 228 MADRONO not occur west of the Invermere area; C. stricta subsp. stricta does not occur in coastal BC. Our study also points to several potential future investigations. A comprehensive DNA investigation would be particularly useful in elucidating the relationships of subspecific enti- ties and species relationships in the C. canadensis, C. stricta, and C. lapponica complex, and could potentially test the new phytogeographical hy- potheses that we have presented, if sufficiently variable DNA markers could be developed. ACKNOWLEDGMENTS Thanks to Andrea to Blasekie for making some of the measurements. 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Fairbarns study site), 5 July 1987, Aiken, S. G. 44 (CAN). British Columbia: Big Bend, logbridge over Wood River where it enters the Columbia at, 100 mi N of Golden, 20 July 1941, Weber, W. A. 2527 (CAN); Toad River, at mi 430 on route 96 between Mucho and Summit Lakes, 28 July 1968, Morton, J. K. NA 1981 (CAN); Mount Robson Provincial Park, Moose Lake, Moose Lake marsh area, 27 Aug 1974, Chuang, Ching Chang 1282 (V); Thompson-Okanagan, Cathedral Provincial Park, Lakeview Mountain, 19 Aug 1980, Douglas, George W., Douglas, Gloria G. 12017 (V); Thompson-Okanagan, Cathedral Provincial Park, km 39 Ashnola Road, 11 June 1980, Douglas, George W., Ratcliffe, Marilyn J., Douglas, G.A. 11771 (V); Liard River Repeater Valley, S aspect, Horseranch area, 21 Aug 1980, Clement, Chris J. HR 8056 (V); Matthew River, flats on E side of, 17 Aug 1982, Pavelick, Leon E. 82 348 (V); Kluskus, NW of Euchu Reach, plot BS-75-106, 27 Aug 1975, Storey, Brenda BS-75-197 (V); Stikine River Kakuchuya Creek, S of, plot LM8025, 6 Aug 1980, Clement, Chris J. LM 80143 (V); Peace River, Flatbed, mi 23.6 on Flatbed- Babcock Murray, plot number AH 76-066, 1976, Harcombe, Andrew AH-139 (V); Cariboo-Chilcotin, Spokin Lake, 2 mi W of the lake, Moffat Creek Road junction, plot BS-76-035, 10 July 1976, Storey, Brenda BS-76-66 (V); Peace River, Fort St. John, Watson Slough, 37 km SW of Fort St. John, 21 Jul 11, Hebda, Richard J., Fitton, Richard 01-61 (V); Peace River, Fort St. John, Watson Slough, 37 km SW of Fort St. John, 21 July 11, Hebda, Richard J., Fitton, Richard 01-64 (V); Peace River, W.A.C. Bennett Dam, 21 July 11, Hebda, Richard J., Fitton, Richard 01-90 (V); Stikine District, Hackett River, at junction of Hackett and Shesley Rivers, 27 July 1971, Schanen, Steven, Schanen, Sara 49,86 (V); Red Pass, 2 mi W of Red Pass Headquarters, 30 June 1974, Chuang, Ching Chang 257 (V); Cheslatta Falls, 2 Aug 23, Richard J. Hebda 00-09 (V); Cheslatta Falls, St. Mary’s Lake, 2 Aug 23, Richard J. Hebda 00- 45 (V); Yukon: Liard River, NW of Watson Lake, 10 Aug 1980, Oswald, E. T. YT-13 (V); Dezadeash River Valley, St. Ellas Mountains, 30 July 1967, Pearson, A. M. 67-313 (CAN); Mayo, 2 Aug 1949, Gillett, J. M., Calder, J. A. 4246 (DAO); Coal River Springs, Coal River Springs proposed park, Site 5, E of Site 4., 6 July 1983, Kennedy, Catherine 34 (DAO); Twin Lakes Campground, Klondike Hwy, km 310, 20 July 1980, Cody, W. J. 28188 (DAO). Calamagrostis canadensis var. langsdor fit: CANADA. British Columbia: Cluculz Lake, E of Vanderhoof, 29 June 1944, Eastham, J. W. 11726 (CAN); Summit Pass, vicinity of Pass, Rocky Mountains, 25 July 1948, Raup, 250 Hugh M., Correll, D. S. 10826 (CAN); Ootsa Lake, Alcan Campground, 2 Aug 24, Richard J. Hebda 00-81 (V); Prince Rupert Forest Region, Telkwa Range, headwaters of Glacis Cr., 2 Sep 1989, Pojar, J. JPS890038 (SMI); Nulki Lake, SW of Vanderhoof, 14 Aug 1945, Munro, J. A. s.n. (V); Van Horlick Creek, near head of E branch, S of Duffy Lake, 4 Sep 1976, Pojar, James J. 760543 (V); Vancouver Island, Enos Lake, along W shore of lake, 8 June 1981, Power, R., Waterhouse, M. 22 (V); Vancouver Island, Haley Lake, Vancouver Island (V); Tatisno Mountain, N of Kitza Lake, 19 Aug 1979, Ceska, Adolf, Ceska, Oldriska, Polster, David F., Martens, Brian 3774 (V); Nadina Lake, Hummock N of Nadina Lake Road, Plot TH-80- 04, 12 Aug 1980, Thompson, C. CT8&0-36 (V); Timothy Mountain, 14 July 1981, Thompson, C. CT&1-112 (V); Cariboo, Big Loon Lake, | mi from the W tip of the lake, 12 July 1976, Storey, Brenda BS-76-101 (V); Far Creek, plot DR#8a, 24 July 1979, Harcombe, Andrew AH-79-11S8 (V); Peace River, Stewart Creek, area on Codner Coldstream, 15 Aug 1979, Ferster, Rick RF-79- 298 (V); Thompson-Okanagan, Hardie Hill, Dewdrop, 21 Sep 1981, Lea, T. T-81-130 (V); Richardson Lake, E.P. 1046, approx. 55 km SW of Burns Lake, Aug 1989, Trowbridge, R., Thomson, S. s.n. (V); Thompson- Okanagan, Ridge Lake, near Kamloops, 31 July 1963, Pringle, William L. s.n. (V); Salahagen Creek, Upper Kimsquit near Salahagen Creek, 23 July 1983, Clement, Chris J. 83132 (V); Flannigan Slough, Taku River area, S end of Flannigan Slough, 10 July 1982, Ceska, Adolf, Ceska, Oldriska, Parisien, L. 12069 (V); Peace River, Watson Slough, 37 km SW of Fort St. John, 21 July 11, Hebda, Richard J., Fitton, Richard 01-63 (V); Gwillum Lake Road , 10 July 1976, C. Clement CJC-70 (V); Haines Road, km 147, near edge of small lake, 19 July 1979, Douglas, George W., Ratcliffe, Marilyn J. 11356 (V); Puggins, Mount, lower end of Puggins Mt. road, 9 July 1979, Pavlick, Leon E., Taylor, B. 79-515 (V); Cumberland Creek, Skeet Club, swamp (V). Yukon: Sheldon Lake, 2 July 19, Cody, W. J., Cody, D. W. 36954 (DAO); Pintail Slough, Old Crow Flats, 12 Aug 1976, Russell, Don 26 (V); Mt. White, Valley slopes and mountain summits about 7 mi E of Little Atlin Lake, 19 Aug 1943, Raup, Hugh M., Correll, D. S. 11434 (CAN); Red Tail Lake, mountain slopes and summits NE of Red Tail Lake, 9 July 1948, Raup, Hugh M., Drury, W. H., Raup, K. A. 13480 (CAN); Canol Rd., km 312, 29 July 1981, Hodgson, Vaughn 433 (DAO); Contact Creek Esso Station, Alaska Hwy, km 949.5, 9 July 1983, Cody, W. J. 32507 (DAO); Tom Creek, area, 13 July 1980, Rosie, R. 951 (V); Tom Creek,, 15 July 1980, Rosie, R. 1410 (V); Kluane National Park, Onion Lake, ca. 46 mi S of Haines Junction, 12 Aug 1973, Douglas, George W., Douglas, Gloria G. 7084 (V); Dry Creek, S of the creek, Mile 1184 Alaska Highway, 27 July 1977, Douglas, Gloria G., Tait, V. L. 10459 (V). RUSSIA: 133827, (V); 133872, 5 Aug 1978, (V); 133861, 24 July 1964, (V); 143959, 26 Aug 1986, (V). Calamagrostis lapponica. CANADA. Alberta: Clear- water Forest Reserve, Baldy Mountain, summit and upper slopes of, N of Nordegg., 18 Aug 1957, Porsild, A. E. 20599, (CAN); Cadomin, S.W.26-45-23-W.S.M., 20 Aug 1967, Pegg, George 2685, (DAO). British Columbia: Birch Mountain, N slope of, Teresa Island, Atlin Lake, 12 Aug 1975, Buttrick, Steven 747, (DAO): Cassiar, Looncry Lake, 2 Sep 1964, Ritcey, Ralph 14, (V); Cassiar, Dall Lake, 13 July 1961, Ahti, Leena, Ahti, Teuvo 6909, (V); Silvertip Mountain, Tootsee Valley, 13 MADRONO [Vol. 58 Aug 1995, Doucet, R., Beaulieu, G. 161, (V); Liard River Basin Petitot River, edge of cutline in moist forest site, 31 July 1974, Haber, Eric, Bergeron, J. 2285, (V); Gladys Lake Ecological Reserve, Ptarmigan Ridge, SE face, 16 July 1975, Pojar, James J. 111d, (V); Stikine District, Gladys Lake Ecological Reserve, Landslide Ridge, Ghost Mountain, 8 July 1975, Pojar, James J. 76e, (V); Gladys Lake Ecological Reserve, Maternity Mountain, 25 July 1975, Pojar, James J. 144g, (V); Stikine River Spatsizi Plateau, above headwaters of Black Fox Creek, 8 Aug 1975, Pojar, James J. 198, (V); Swift River, 10 km SW of, 28 July 1980, Brayshaw, T. Christopher 80-160, (V); Horseranch, N end of Horse- ranch, 24 Aug 1980, Clement, Chris J. HR 8094, (V); Horseranch Lake, N end of, 24 Aug 1980, Clement, Chris J. HR8095, (V); Garbutt Creek, Garbutt Creek area, 25 Aug 1979, Ceska, Adolf; Ceska, Oldriska, Polster, David F., Martens, Brian 3562, (V). Northwest Territories: MacKenzie district, Eskimo Lake Basin, outlet of Sitidgi Lake, 11 Aug 1957, Cody, W. J, Ferguson, D. H. 10815, (SMI). Yukon: Ptarmigan Heart, mountain slopes and summits NE of Ptarmigan Heart, 13 July 1948, Raup, Hugh M., Drury, W. H., Raup, K. A. 13618, (CAN); Tatshenshini River, mi 100, Haines Highway, 14 Aug 1957, Schofield, W. B., Crum, H. A. 8265, (CAN); Little Atlin Lake, Valley slopes and mountain summits, 13 Aug 1943, Raup, Hugh M., Correll, D. S. 11287, (CAN); Ptarmigan Heart, mountain slopes and summits NE of Ptarmigan Heart, 16 July 1948, Raup, Hugh M., Drury, W. H., Raup, K. A. 13720, (CAN); LaBiche River area, Kotaneelee Range, 20 June 1998, Rosie, Rhonda 2069, (DAO); Otter Lake, above lake S of Itsei Range, 2 Aug 1960, Calder, J. A., Kukkonen, I. 27759, (DAO); Kusawa Lake, mountain between Kusawa Lake and Jojo Lake, 19 Sep 1997, Bennett, B. 97-672, (DAO); Deep Creek, about 2 mi from the Arctic Ocean, 16 Aug 1976, Russell, Don 4, (V); Ferry Hill, 9 Aug 1977, Rosie, Rhonda 474, (V); Mt. Laborite, 20 July 1994, Zoladeski, Chris 200, (DAO). USA. Alaska: Nabesna Rd, mi 89, 24 July 1947, Dutilly, LePage and O'Neill 21563, (DAO); Kurupa Valley, western side of Kurupa Valley about 8 mi NW of Kurupa Lake, 3 Aug 1952, Riedeman, Robert s.n., (DAO); King Salmon, 8 Aug 1952, Schofield, W. B. 2653, (DAO); Eastern Brooks Range, Porcupine Lake, 24 July 1979, Gustafson, Karen s.n. (V). Calamagrostis montanensis. CANADA. Alberta: Liv- ingston Valley, at the gap, 11 Aug 1915, Malte, M. O 108222 (DAO); Cowley, SE of, 9 Aug 1939, Moss, E. H. 345 (DAO); Carway, at U.S. border, 15 mi S of Cardston, 2 Aug 1950, Dore, W. G. 12269 (DAO); Wapato River, Wembley Region, S of Wembley, 8 July 1976, Barkworth, M. 1459 (DAO); Tough Creek, extreme SW corner, sect 17, S of, 28 July 1982, Aiken, S. G., Darbyshire, S. J., Klumph, Bud 2506 (DAO). British Columbia: Invermere, Old Fort Community Hall, Columbia Valley, 16 July 1947, Eastham, J. W. 15930 (DAO); Windermere Beach, 28 July 1947, Eastham, J. W. 15920 (DAO); Invermere, Old Fort Community Hall, Columbia Valley, 16 July 1947, Eastham, J. W. 15930 (DAO); Kootenay, Invermere, 11 Aug 1943, Eastham, John W. 11106 (V); Kootenay, Edgewater, between Sinclair Creek and Edgewater, Columbia Valley, 29 July 1947, Eastham, John W. s.n. (V); Kootenay, Invermere, Old Fort Community Hall, 16 July 1947, Eastham, John W. s.n. (V); Kootenay, Windermere Lake, Columbia Valley, above Wind- 2011] ermere Beach, 28 July 1947, Eastham, John W. 20.949 (V); Kootenay, Fairmont Hot Springs, one mi N of, 2 July 1948, Eastham, John W. s.n. (V); Peace River, Clayhurst Ecological Reserve, Doe Creek, 29 July 1969, Brayshaw, T. Christopher 5352 (V); Kootenay, Inver- mere, | July 1947, Fodor, Fred s.n. (V); Kootenay, Wilner Marsh, slope above, Plot 6102-01, 13 June 1981, Lea, T. TL-81-06 (V); Kootenay, Radium, N-most highway viewpoint S of Radium, 1.7 km S of main intersection of Radium, 25 Aug 1995, Roemer, Hans L. 95077 (V); Alces River, 26 July 1995, Douglas, George W., Djan-Chekar, Nathalie 13060 (V). Calamagrostis nutkaensis. CANADA. British Colum- bia: Mayer Lake, S end of lake, W of Tlell, Graham Island, 20 Aug 1964, Calder, J. A., Taylor, R. L. 36105 (DAO); Prince Rupert, 30 July 1916, Malte, M. O. 106868 (DAO); Henslung Bay, near Bay, Langara Island off NW tip of Graham Island, 16 July 1957, Calder, J. A., Savile, D. B. O., Taylor, R. L. 22534 (DAO); Vancouver Island, Kyuquot, Markale, 3 Aug 1957, Bell, Marcus, Davidson, John 767 (V); Queen Charlotte Islands, Tow Hill, Graham Island, 30 May 1963, Young, A., Hubbard, W. 110 (V); North Coast, Digby Island, Prince Rupert Airport, 9 Aug 1973, Brayshaw, T. Christopher s.n. (V); Vancouver Island, Cape Scott, near lighthouse, 22 July 1982, Pavlick, Leon E. 82-60 (V); Queen Charlotte Islands, Rennell Sound, Re9B4, 27 July 1982, Ryan, Michael W. 2 (V); Brooks Peninsula, Cladothamnus Lake, Ridge Quadrant, Ridge SW above lake, 9 Aug 1981, Ogilvie, Robert T., Hebda, Richard J., Roemer, Hans L. 81893 (V); Gillen Harbour, head of Harbour, Dewdney Island, 13 July 1984, Ogilvie, Robert T., Roemer, Hans L. 8471318 (V); Dewdney Island, SW peninsula of, 13 July 1984, Ogilvie, Robert T., Roemer, Hans L. 8471354 (V); Dewdney Island, head of Gillen Harbour, 13 July 1984, Ogilvie, Robert T., Roemer, Hans L. 8471319 (V); Brooks Peninsulal900 Peak, Lagoon Quadrant, 1900 Peak, 9 Aug 1984, Ogilvie, Robert T., Schofield, Wilfred J., Hebda, Richard J. 848932 (V); Vancouver Island, Pacific Rim National Park, Nettle Island, near camp, Barkley Sound, Broken Islands, 8 Aug 1982, Ogilvie, Robert T., Hebda, Richard J. 828089 (V); Central Coast, Dundas Islands, Zayas Island, 25 July 1987, Ceska, Adolf, Ceska, Oldriska, Ogilvie, Robert T. 22032 (V); Vancouver Island, Bamfield, SW of Bamfield, 22 Aug 1980, Ogilvie, Robert T. s.n. (V); Queen Charlotte Islands, Geikie Creek, Graham Island, 200 m S of Geikie creek #3, 50 m E of highway 16, 22 Aug 1997, Lomer, Frank, Grove, N. 97529 (V); Gulf Islands, Egeria Mountain, Southern Bowl, Porcher Island, 8 Aug 1987, Ceska, Adolf, Ceska, Oldriska 22649 (V); Central Coast, Dundas Islands, Zayas Island, 25 July 1987, Ceska, Adolf, Ceska, Oldriska, Ogilvie, Robert T. 22053 (V): Central Coast, Campbell Island, N of Bella Bella (Waglisla), Wag air fueling station, 19 July 1999, Hebda, Richard J. s.n. (V). Calamagrostis purpurascens. CANADA. Alberta: Jasper National Park, Mt. Edith Cavell, 28 Aug 1964, Calder, J. A 37200 (DAO); Jasper National Park, Lake Edith, YMCA Lodge, 4 July 1955, Jenkins, L. 5817 (DAO). British Columbia: Cariboo, Mt. Begbie, 23 June 1944, Eastham, John W. 17026 (V); Cariboo-Chilcotin, Sinkut Mountain, near Vanderhoof, 18 July 1945, Eastham, John W. 18828 (V); Cassiar District, Cassiar, 18 June 1956, Taylor, Thomas M.C., Szczawinski, Adam F., Bell, Marcus 391 (V); Cariboo, Mt. Pope, a few mi NW of Fort St. James, 11 July 1892, Hatcher, J. s.n. MARR ET AL.: MORPHOLOGY OF CALAMAGROSTIS IN B.C., CANADA 231 (V); Liard River, Liard Hot Springs Park, 16 Aug 1971, Brayshaw, T. Christopher, Barrett, David s.n. (V); Liard River, Liard Hot Springs Provincial Park, Mt. Ole, 7 July 1971, Brayshaw, T. Christopher, Barrett, David s.n. (V); Cariboo-Chilcotin, Taseko River, valley side on E by Taseko River Rd, 14 July 1978, Pavlick, Leon E., Sax, Michael 78-571 (V); Hutchison Lake, near lakes, 21 Aug 1979, Ceska, Adolf, Ceska, Oldriska, Polster, David F., Martens, Brian SO6I (V); Buckley Creek, above Klastline River, upstream Buckley Creek, 29 Aug 1979, Ceska, Adolf, Ceska, Oldriska, Polster, David F. 8073 (V); Kootenay, Flathead, two ridge tops in S, ca. 5 mi from Canada-US border, July 1973, Dick, John FR24 (V); Kootenay, Skookumchuck, 0.1 mi up Regional Garbage Dump Rd, 2 July 1976, Ferster, Rick 76-78 (V); Dean River, Upper Dean River Rd, plot number 2803-79, 1979, Harcombe, Andrew AH-79- 30 (V); Stikine River, Telegraph Creek, slopes above Day’s Ranch, 10 June 1980, Ceska, Adolf, Ceska, Oldriska, Polster, David F. 4070 (V); Cariboo-Chilcotin, Cheslatta, Kritchlow property, | km E of Cheslatta, 21 July 12, Hebda, Richard J., Fitton, Richard 01-119 (V); Tanzilla River above the river, 2 km above junction with Stikine River, 10 July 1980, Ceska, Adolf, Ceska, Oldriska, Polster, David F. 8054 (V). USA. Alaska: Talkeetna Mts., 2 Sep 1978, Talbot, S. S. T8023-V-15 (DAO). Washington: Buckhorn Mt., T27N R4W S13 SE1/4, 8 June 1979, Buckingham, Nelsa 2129 (OLYM); Olympic National Park, Royal Basin, RB2, Ridge between Royal Creek and Dungeness River, Aug 1983, Dalton, Burger 2656 (OLYM). Calamagrostis rubescens. CANADA. British Colum- bia: Bear Cr. Falls, N of Clearwater station, Bear Cr. Falls, 10 Aug 1956, Calder, J. A., Parmelee, J. A. Taylor, R. L. 19916 (DAO); Invermere, 31 July 1915, Malte, M. O. 108288 (DAO); Quesnell, S. of Quesnell, 14 July 1982, Aiken, S. G., Darbyshire, S. J., Roberts, A. 2320 (DAO); Alexis Creek, 16 July 1982, Aiken, S. G, Darbyshire, S. J. 2365 (DAO); Saxton Lake, 21 July 2008, Richard J. Hebda, Richard Fitton OO-37 (V); Cariboo-Chilcotin, Vanderhoof, 15 Aug 1919, Macoun, John M. 27 (V); Kootenay, Cranbrook, 7 Aug 1943, Eastham, John W. 16385 (V); Cariboo-Chilcotin, Riske Creek, N slope, SW of Beecher Dam, 3 Oct 1968, Brayshaw, T. Christopher s.n. (V); Thompson-Okanagan, Manning Provincial Park, Wrangler Station, approx- imately one half mi from Nature House, 16 Aug 1973, Chuang, Ching Chang 1259 (V); Lower Fraser Valley, Ross Lake, | mi N of International Boundary on Ross Lake Rd, 20 July 1971, Smith, R. B. 29 (V); Kootenay, Arrow Lakes, area, 12 June 1975, Polster, David s.n. (V); Kootenay, Akamina Creek, 0.8 km downstream from Gloyne Camp on N side of road, 21 Aug 1975, Polster, Alan, Plug, Egbert, Polster, David 80:14 (V); Cariboo- Chilcotin, Riske Creek, south creek area, 11 July 1978, Pavlick, Leon E., Sax, Michael 78-452 (V); Cariboo- Chilcotin, Dragonfly Lake, E of Dragonfly Lake, near Deka Lake, 13 July 1972, Resource Analysis Branch, Kelowna 72-92 (V); Kootenay, Argenta, Johnson’s Landing, 6 mi S of Argenta-Johnson’s Landing inter- section, 15 June 1982, Wood, Terry 822085 (V); Kootenay, Harmer Ridge, Natal, 30 July 1973, Dick, John HR 9 (V); Thompson-Okanagan, Peachland Creek, Peachland Creek area, 8 July 1987, Pavlick, Leon E. 87-282 (V); Cariboo-Chilcotin, Dragonfly Lake, E of Dragonfly Lake, near Deka Lake, 13 July 1972, van Barneveld, Jim W. JvB-72-92 (V); Cariboo-Chilcotin, 232 Oregon Jack Creek, 5 July 1978, Pavlick, Leon E., Sax, Michael 78-271 (V). Calamagrostis sesquiflora. CANADA. British Colum- bia: Queen Charlotte Islands, Bigsby Inlet, head of Bigsby Inlet, opposite Lyell Island, E coast of Moresby Isl., 5 July 1957, Calder, J. A., Taylor, R. L., Saville, D. B. O. 22141 (DAO); Queen Charlotte Islands, Mt. de la Touche, Farifax Inlet, Tasu Sound, west coast of Moresby Isl., 16 Aug 1957, Calder, J. A., Taylor, R. L. 23571 (DAO); Queen Charlotte Islands, Mosquito Lake, Mt. above Mosquito Lake near head of Cumshewa Inlet, 24 Aug 1957, Calder, J. A., Taylor, R. L. 23753 (DAO); Queen Charlotte Islands, Victoria Lk., Upper Victoria Lk., neaer S end Moresby Isl, 5 July 1964, Calder, J. A., Taylor, R. L. 35718 (DAO); Queen Charlotte Islands, Cumshewa Inlet, 3 mi W of head of Cumshewa Inlet below N face of Mt. Moresby, 1 Aug 1964, Calder, J. A., Taylor, R. L. 36507 (DAO); Queen Charlotte Islands, Graham Island, E side of Shields Bay, Rennel Sound, W coast of the island, 16 July 1963, Brassard, Hainault 2824 (V); Brooks Peninsula, Ridge Quadrant, Ridge SW above Cla- dothamnus Lake, 9 Aug 1981, R. 7. Ogilvie, R.J. Hebda & Hans L. Roemer 81894 (V); Vancouver Island, Brooks Peninsula, Cassiope Pond, ridge quadrant, crest of ridge E of the pond, 31 July 1981, Ogilvie, Robert T., Hebda, Richard J., Roemer, Hans L. 8173113 (V); Vancouver Island, Doom Mt., summit of main peak, ridge quadrant, Brooks Peninsula, 17 Aug 1981, Ogilvie, Robert T., Hebda, Richard J., Roemer, Hans L. 8181711 (V); Vancouver Island, Brooks Peninsula, July 1978, Roemer, Hans L. 7890 (V); Queen Charlotte Islands, Chanal, Port, W Graham Island, July 1979, Roemer, Hans L. 79159 (V); Queen Charlotte Islands, Takakia Lake, North Ridge, 19 July 1980, Ogilvie, Robert T., Roemer, Hans L., Mersereau, W.O. s.n. (V); Anna Lake, North end, upper waterfall, 25 Aug 1992, Ogilvie, Robert T. s.n. (V); Queen Charlotte Islands, Mount Laysen, 20 Aug 1992, Ogilvie, Robert T. s.n. (V); Queen Charlotte Islands, Dinan Creek, on mountain ridge, at headwaters of Dinan creek, Graham Island, 17 July 1997, Lomer, Frank, Grove, N. 97386 (V); Queen Charlotte Islands, Apex Mt., 3 km W of Apex Mt., Moresby Island, 19 Aug 1997, Lomer, Frank, Grove, N. 97448 (V); Queen Charlotte Islands, Moresby Island, Mosquito Mt. , 25 km SSW from Queen Charlotte City, 20 Aug 1997, Lomer, Frank, Grove, N. 97498 (V). Calamagrostis stricta subsp. inexpansa. CANADA. Alberta: Beaverlodge, 17 July 1921, Malte, M. O. 106930 (DAO); Willow Creek, Willow Cr. area, Jasper NP, 1978, Reynolds, H. J75 (DAO). British Columbia: Noralee, Francois Lake, above Brewer’s, 8 July 1944, Eastham, J. W. 11866 (CAN); Buckinghorse River, 31 Aug 1943, Raup, Hugh M., Correll, D. S. 11592 (CAN); Queen Charlotte Islands, Delkatla Inlet, just E of Masset, Graham Island, 3 Oct 1968, Brayshaw,_ T. Christopher s.n. (V); Peace River, Portage Mt., Portage Mt. Dam, small lake beside the BC Hydro camp, 25 July 1965, Szezawinski, Adam F. 8/65 (V); Alsek River, Tatshenshini River, junction of, 24 June 1975, Brayshaw, T. Christopher, Carriagan, C. J. s.n. (V); Kootenay, Mt. Robson Provincial Park, S of Nature House along Fraser River, 22 July 1975, Chuang, Ching Chang 75/75 (V); Vancouver Island, McCreight Lake, 7 Oct 1977, Brayshaw, T. Christopher s.n. (V); Pacific Rim National Park, Klanawa River, 11 July 1973, Hartwell, Sharon 71105 (V); Vancouver Island, Wickaninnish Bay, along Long Beach near the mouth of Sandhill MADRONO [Vol. 58 Creek, 20 June 1969, Soper, James H., Brayshaw, T. Christopher, Shchepanek, Michael J. 12307 (V); Bea- verdam Lake, 18 Aug 1978, Brayshaw, T. Christopher 78-677 (WV); Stikine District, Kuachon Lake, NE of Kuachon Lake Lodge, 29 Aug 1979, Ceska, Adolf, Ceska, Oldriska, Polster, David F. 4163 (V); Narraway River, E of Manitou Mtn., 18 Aug 1977, Ceska, Adolf, Wood, Terry 9289 (V); Kootenay, Tete Jaune Cache, on Highway 5 near, 8 July 1977, Ceska, Adolf, Wood, Terry 9290 (V); Vancouver Island, Farewell Lake, N of Campbell River, 16 Aug 1983, Ceska, Adolf, Ceska, Oldriska 16283 (V); Vancouver Island, Pacific Rim National Park, Effingham Island, Barkley Sound, Broken Islands Group, 8 Aug 1982, Ogilvie, Robert T., Hebda, Richard J. 828083 (V); Thompson-Okana- gan, Minnie Lake, by pothole lake just N of, 5 July 1987, Pavlick, Leon E. 87-168 (V); Kootenay, Ewin Creek, 0.8 mi SW of Ewin Creek on Main Fording Coal Rd, 9 Sept 1977, Ferster, Rick 77-98 (V); Vancouver Island, Keeha Beach, delta of Keeha Creek, 23 June 1983, Ogilvie, Robert T. s.n. (V); Gulf Islands, Trial Island, 20 July 1976, Ceska, Adolf; Ceska, Oldriska s.n. (V); Atlin, around Tarahne steamboat and Atlin Inn, 15 July 1982, Ceska, Adolf; Ceska, Oldriska, Goward, Trevor 12649 (V); One Fifteen Creek, mi 406, 3 km W of 115 Creek picnic site, 26 July 1982, Ceska, Adolf, Ceska, Oldriska, Goward, Trevor 13469 (V); Vern Ritchie Glacier, foreland of glacier, Haines Triangle, 26 July 1992, Pojar, James J. JP920156 (V); Peace River, Cecil Lake, off Road #245, enclosures near lake, 25 June 1997, Ceska, Adolf 30730 (V). Yukon: Canol Rd., lower part of Canol road, along the road, 13 July 1947, Porsild, M. P., Porsild, R. T. 506 (CAN); Hunker P.O., 8 July 1950, Campbell, John D. 50 (DAO); Eagle Plains, E of the Richardson Mtns., July 1979, James, T. D. W. 17 (DAO); Haines Junction, on Haines High- way S of Haines Junction, 26 July 1980, Cody, W. J, Ginns, J. H. 28384 (DAO); Hyland River, 25 July 1994, Brunner, Greg 84 (DAO); Itsi Range, unnamed lake in range near Yukon-McKenzie border, 31 July 1960, Calder, J. A., Kukkonen, I. 27642 (DAO); Klondike Highway, km 656, 19 July 1980, Cody, W. J. 28090 (DAO); Canol Rd, 10 km, 1 Aug 1980, Cody, W. J., Ginns, J. H. 28767 (DAO); Dawson, along bank of Yukon River at Dawson., 18 July 1930, W. J. G. s.n. (V). Calamagrostis stricta subsp. stricta. CANADA. Alberta: Pigeon Lk., 15 Aug 1945, Turner, G. H. 4680 (DAO); Pigeon Lake, Opposite Ma-Me-O Beach on Pigeon Lk., 29 July 1947, Turner, G. H. 5927 (DAO); Falher, 19 July 1948, Jenkins, L. 566 (DAO); Many- berries, 17 June 1937, Campbell, J. A. s.n. (DAO). British Columbia: Kleena Kleene, 3.5 mi W of Kleena Kleene P.O., 5 July 1956, Calder, J. A., Parmelee, J. A. Taylor, R. L. 19196 (DAO); Cariboo-Chilcotin, Burns Lake, swamp at Tatlarose S of lake, 9 July 1944, Eastham, John W. 17759 (V); Stikine District, Lake Tatogga, NE end of, 10 Aug 1971, Brayshaw,_ T.: Christopher, Barrett, David s.n. (V); Barney Lake, mi 581, Alaska Hwy, | Aug 1974, Brayshaw, T. Christo- pher, Polster, David F. s.n. (V); Peace River, Stony Lake, 17 mi N of, Peace River, district, 5 Aug 1976, Chuang, Ching Chang 493 (V); Fletcher Lake, 3.2 km NE of, 5 Aug 1978, Thompson, Carol E. s.n. (V); Peace River, Bear Flat, Bear Flat area, 19 July 1979, Pavlick, Leon E., Taylor, B. -797 (V); Cariboo-Chilcotin, Hooch Lake, S of, ca. 15 km SW of Nimpo Lake, 7 July 1980, Annas, Richard, Ruyle, Gloria G., Nicholson, Allison, 2011] Coupe, Ray 80-272 (V); Liard River Basin, Coal River, N of, 20 Aug 1979, Ceska, Adolf, Ceska, Oldriska, Polster, David F., Martens, Brian 3726a (V); Upper Dean River, plot number PR#4, 16 July 1979, Harcombe, Andrew AH-79-40 (V); Cariboo-Chilcotin, Lessard Lake, N of the lake, plot no. Dr#7, 22 July 1979, Harcombe, Andrew AH-79-101 (V); Peace River, MARR ET AL.: MORPHOLOGY OF CALAMAGROSTIS IN B.C., CANADA 205 Watson Slough, 37 km SW of Fort St. John, 21 July 11, Hebda, Richard J., Fitton, Richard 01-56 (V); Cheslatta Lake, floodplain, 2 Aug 23, Richard J. Hebda 00-21 (V). Yukon: Kluane National Park, Haines Junction, one mi N of the junction of Kawkawulsh and Dezadeash Rivers and ca. 10 mi WSW of Haines Junction, | Aug 1973, Douglas, G. W., Douglas, G. G. 6682 (CAN). MADRONO, Vol. 58, No. 4, pp. 234-248, 2011 POPULATION ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER (PINUS BALFOURIANA) WITH A DISJUNCT DISTRIBUTION IN CALIFORNIA PATRICIA E. MALONEY Department of Plant Pathology & Tahoe Environmental Research Center, University of California, One Shields Ave, Davis, CA 95616 pemaloney@ucdavis.edu ABSTRACT Foxtail pine, Pinus balfouriana Grev. & Balf., is an endemic subalpine conifer of California with two allopatric subspecies. The northern subspecies grows in northwestern California and the southern subspecies is found in the southern Sierra Nevada. We studied three northern and three southern populations of P. balfouriana to evaluate the population biology, demography, mortality agents, and environmental conditions in these contrasting regions. Northern populations exist under a mesic climatic regime, a diversity of geological substrates, stands of high tree species richness, lower densities and basal areas, but higher numbers of recruitment, with a relatively mixed size class distribution. Southern foxtail pine populations exist under a xeric climatic regime dominated by granitic substrates, with moderate to high stand densities, less tree species-rich, lower recruitment numbers, but higher basal areas; due to a skewed size class distribution with high representation of large diameter trees. Recruitment in the north averaged 169.3 seedlings/saplings ha~', compared to 91.3 seedlings/saplings ha~' in the south, despite the fact that northern populations produce less cones on average (3695 cones ha~') than populations in the southern Sierra Nevada (7642 cones ha™'). At the stand-level, solar radiation input and foxtail pine density were correlated with fecundity. These factors may correspond with microenvironmental and topographic conditions that favor germination (e.g., warmer microclimate) and propagule pressure (e.g., seed supply). At the local or plot-level, microenviron- mental conditions (e.g., litter, substrate type, and microhabitat) and factors corresponding to local seed supply (e.g., density, basal area, number of cones, and number of reproductive adults) were correlated with recruitment, particularly in the southern Sierra Nevada. Foxtail pine is recruiting episodically in higher numbers in the north and lower numbers in the south. Four of six populations appear to be stable, due to low mortality and high survivorship. Low estimates of population growth (A) at Lake Mountain (north) and Sirretta Peak (south) were due to mortality of large diameter trees and low recruitment. At these locations, mountain pine beetle-mediated mortality and drought stress appear to be important factors contributing to current population trends. Key Words: Cronartium ribicola, demography, endemic conifer, Foxtail pine, Pinus balfouriana, population structure, recruitment, subalpine. Species and ecosystems at high latitudes and elevations are considered to be the most sensitive to global climate change (Parmesan 2006). A recent bioclimatic model has predicted significant range contractions for an important subalpine conifer (whitebark pine, Pinus albicaulis Engelm.) in western North America (Rehfeldt et al. 2006). This model and others like it, that project species distributions, are based on climate variables and species abundance (e.g., Lenihan et al. 2003; Rehfeldt et al. 2006; Beaumont et al. 2007). However, these models lack key information on population dynamics (e.g., fecundity, survival, mortality, and growth), dispersal, biotic interac- tions (e.g., competition, disease, and insects), genetics, and environmental heterogeneity. Clark et al. (2011) highlight the importance of incor- porating demographic parameters (fecundity, mortality, survival, growth) in evaluating species responses to climate change. With that said, field- based ecological and demographic approaches are necessary to understand basic population biology and in turn the potential vulnerability of forest tree species to natural and anthropogenic disturbances. Non-native pathogens, climate change, and climate-driven outbreaks of native insects are three threats to high elevation white pines in western North America (Tomback and Achuff 2010). Foxtail pine (Pinus balfouriana Grev. & Balf. subsp. ba/fouriana) is a high elevation white pine endemic to California, with two allopatric subspecies that are separated by approximately 500 km; one in the Klamath, Scott, and Yolla Bolly Mountains of northwestern California and the other in the southern Sierra Nevada (Mas- trogiuseppe and Mastrogiuseppe 1980; Oline et al. 2000; see Fig. 1). For some time, the range disjunction was thought to have occurred during the Holocene Xerotherm (4000-8000 years ago; — Axelrod 1976, 1977). Using a contemporary © molecular genetic approach, Eckert et al. (2008) © propose that this range disjunction occurred © much earlier; in the Middle (0.13—0.86 million 2011] MALONEY: ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER — 235 0 50 Lae 100 Kilometers 0 50 100 Miles aes Has FIG. 1. source: Little (1999). years ago [Mal]) to Early Pleistocene (0.93— 2.45 Ma). This time period corresponds with the Sherwin glaciation (~1 Ma) in the Sierra Nevada, one of the largest glacial episodes during the Pleistocene (Hill 2006; Eckert et al. 2008). Geographic and temporal separation of the two subspecies corresponds with significant genetic, morphological, biochemical, and envi- ronmental differences (Bailey 1970; Snajberk et al. 1979; Mastrogiuseppe and Mastrogiuseppe 1980; Oline et al. 2000; Eckert et al. 2008; Eckert et al. 2010); potentially influencing the popula- tion biology and demography of foxtail pine in the two regions. For example, the northern subspecies (P. b. balfouriana) grows between 2050 and 2600 m, generally on the highest peaks Q&S Pinus balfouriana demographic plots [ Pinus balfouriana distribution Location of study areas and foxtail pine distribution in California. Pinus balfouriana distribution map and ridges, and separated by deep valleys (Oline et al. 2000; Eckert et al. 2010). The “mountain island” nature of foxtail pine populations in the north results in little gene flow and large genetic diversity among stands (Oline et al. 2000; Eckert et al. 2010). Foxtail pine is often a dominant component in these stands but mixes with a diversity of subalpine conifer species such as red fir (Abies magnifica A. Murr.), mountain hem- lock (Tsuga mertensiana [Bong]. Carriere), lodge- pole (P. contorta Douglas ex Loudon), whitebark (P. albicaulis), western white (P. monticola Douglas ex D. Don), and Jeffrey (P. jeffreyi Grev. & Balf.) pines (Oline et al. 2000; Eckert 2006). In this region the maximum attainable age for foxtail pine has been estimated to be between 236 1300-1500 years (Mastrogiuseppe and Mastro- giuseppe 1980). The southern subspecies (P. b. austrina R. J. Mastrog. & J. D. Mastrog.) in the southern Sierra Nevada grows between elevations of 2700 and 3600 m and defines timberline in this region. Here, stands are generally larger and more contiguous with less genetic differences among them, relative to differences within stands (Oline et al. 2000). In the south, foxtail may grow in pure stands or in association with species such as lodgepole, limber (P. flexilis E. James), or whitebark pine. Maximum attainable age for foxtail pine in the southern Sierra Nevada has been estimated to be between 2500-3000 years (Mastrogiuseppe and Mastrogiuseppe 1980). Mountain climates throughout the range of foxtail pine are largely dominated by the Califor- nia Mediterranean climatic regime, characterized by cold, wet winters, with long, warm, and dry summers. Within the Mediterranean climatic parameters, precipitation totals and growing season lengths vary considerably. The mountains of interior northwestern California receive high amounts of rainfall (=1000—1500 mm) compared to the southern Sierra Nevada (<1000 mm). January minimum and July maximum tempera- tures differ considerably between the two regions as well, with warmer temperatures and longer growing seasons in the north compared to the south. In the north, foxtail occurs on a diversity of geological substrates and in the southern Sierra Nevada foxtail pine almost exclusively grows on granitic substrates (USDA, NRCS 2008). Historical stand dynamics (e.g., mortality and recruitment) have been inferred from dendro- chronological studies of foxtail pine in the southern Sierra Nevada (Lloyd 1997; Lloyd and Graumlich 1997). These studies have shown that treeline populations of foxtail pine have fluctu- ated in elevation in response to changes in both temperature and precipitation (Scuderi 1987; Lloyd 1997; Lloyd and Graumlich 1997). In the Klamath Mountains environmental heterogene- ity (e.g., substrate type, microsite conditions, topography, and species composition) can strong- ly influence not only the persistence of foxtail pine but also recruitment success and subsequent downslope expansion (Eckert 2006; Eckert and Eckert 2007). In a recent study Crimmins et al. (2011) report downhill shifts in numerous plant species in California; largely tracking climatic water balance rather than temperature. In the southern Sierra Nevada, recruitment success appears to be influenced by soil moisture and topographic position (e.g., slopes with higher radiant input), with recruitment patterns being episodic (Bunn et al. 2005). Given autoecological and genetic differences between the two subspecies the objectives of the study were to determine: (7) population and stand characteristics of foxtail pine in the regions of the MADRONO [Vol. 58 northern and southern subspecies, (ii) factors important to recruitment, and (ii/) current structure and population trends. Knowledge of the population biology, demographics (e.g., survival, fecundity, and growth), and environ- ment of foxtail pine populations in both regions is central to understanding how vulnerable this endemic conifer is to natural and anthropogenic disturbances (Cronartium ribicola J. C. Fisch.— cause of white pine blister rust [WPBR], out- breaks by the native insect Dendroctonus ponder- osae Hopkins [mountain pine beetle, MPB], climate change, and fire) and how these might influence future populations of this narrowly distributed white pine. MATERIALS AND METHODS Study Sites During the summers of 2008—2009, we select- ed six study populations, with two or three permanent demographic plots per population (sampling area within a population = 4 ha), for a total of 16 plots on National Forest System lands in California (Fig. 1). Only two demo- graphic plots were established at both North Yolla Bolly and Sirretta Peak due to small population sizes and logistics. Each of the six populations was located in a distinct watershed and distributed in the northern (three sites) and southern (three sites) regions to capture varia- tion in the physical environment (e.g., climate, geology, topography, forest composition; see Fig. 1). Within the northern and southern sub- species distribution, study sites were located in the northern, central, and southern portions of foxtail pine’s geographic range in each of those regions (Fig. 1). In northwestern California, study populations were located at Lake Moun- tain, Mount Eddy, and the North Yolla Bolly (listed from north to south). In the southern Sierra Nevada, study populations were located at Onion Valley, Cottonwood Pass, and Sirretta Peak (north to south, respectively). Population and Stand Sampling Once a population was located, a random starting point was chosen for the first plot; the second and third plots were sited =100 m from the first plot. Within a population, each of three replicate plots were 40 m X 100 m (4000 m7’) with sampling covering approximately 1.2 hectares within a 4-hectare area. The following data were recorded for each demographic plot: GPS loca- _ tion (UTM: NAD27 coordinates), slope (in percent), aspect, elevation (in meters), visible signs of past fire or ignition (i.e., basal fire scar, — bole scorch, lightning strike), slope position (ridge-top, upper slope, mid-slope, lower slope, 2011] MALONEY: ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER — 237 valley bottom, or bench), and land-use history (e.g., historical logging, fire suppression, recent thinning, use of wildland fire, recreation, none), site condition (e.g., xeric or mesic), presence or absence of the Clark’s nutcracker (Nucifraga columbiana). Clark’s nutcracker is an important dispersal agent of many western pine species including the high elevation white pines such as whitebark pine, limber pine, and Great Basin bristlecone pine, P. /ongaeva D.K. Bailey (Lanner 1982, 1988: Tomback and Linhart 1990). Within each demographic plot, all P. balfouri- ana were identified and diameter at breast height (d.b.h. in cm) recorded for all individual stems =1.37 m tall. Seedlings and saplings were all stems <1.37 m in height. All tree positions (x and y coordinates from the centerline of the plot) were recorded and mapped and data collected for tree status (live or dead), crown condition (rating 1-10 as follows: 1: =10% dead, dying, damaged, infected; 2: 11-20% dead, dying, damaged, infected; 3: 21-30% dead, dying, infected, etc.), and crown position (understory, suppressed, intermediate, codominant, dominant, or open). Signs and symptoms of pathogens (e.g., WPBR, dwarf mistletoe, and root diseases) and insects were also recorded. Dendroctonus ponderosae (MPB) was confirmed if there was the presence of pitch tubes, frass, and characteristic galleries (Furniss and Carolin 1977). Reproductive output was assessed by counting the number of current and previous years’ cones per tree. Seedlings and saplings <1.37 m tall were evaluated within each demographic plot by establishing three nested recruitment subplots that were 15 m X 15 m in size (totaling 225 m‘’), for a total of nine regeneration plots/population (six regeneration plots for North Yolla Bolly and Sirretta Peak). All recruitment was counted and identified to species. For P. balfouriana recruit- ment, data were collected on basal diameter (cm), height (cm), crown condition, status (live or dead), disease condition, and whorls counted for aging. Microenvironmental conditions for each foxtail recruit were evaluated by measuring litter depth (cm), substrate type (e.g., exposed soil, decomposed granite, soil and litter, log, rock), canopy type (open/exposed, partially closed, and closed), and microhabitat condition (tree/shrub/ log nurse, rock shelter, other, or none). A limited number of foxtail seedlings/saplings were sam- pled to obtain size-age relationships by counting growth rings, as well as measuring height, diameter, and number of whorls. We used a multiple linear regression to estimate age for recruitment present in demographic plots. Inde- pendent variables loaded into the regression model were height, whorl count, and diameter of field-sampled foxtails. Diameter explained 97% of the variation in the model and yielded parameter estimates and the regression equation Y = 22.2949X, + 0.7683; r?7 = 0.971 (Fig =169.19, P < 0.0001). This equation was then used to estimate age and the year that a seedling or sapling had been recruited into a demographic plot. A forest vegetation plot (40 m x 40 m) was nested within each demographic plot to obtain tree data for other tree species besides P. balfouriana (1.e., d.b.h., status, diseases, insects, crown condition, crown class, etc.). All tree and recruitment data at each plot were collected to quantify stand structure, composition, basal area, and density. Positions of all associate trees were recorded and mapped. For each demographic plot, climatic parame- ters of mean, minimum, and maximum monthly and annual temperature and precipitation from the period of 1971-2000 were provided by FHTET (USDA FS Forest Health Technology Enterprise Team, Fort Collins, CO) using the PRISM climatic model (Daly et al. 1994). Parent material and soil survey data were provided by the South Lake Tahoe office of the USDA Natural Resources Conservation Service (NRCS). Percentage maximum solar radiation input was calculated using slope and aspect (Buffo et al. 1972). A check of collinearity in the multiple linear regression model, to estimate recruitment age, was done employing leverage plots and bivariate scatterplots. Assumptions of normality and homogeneity of variances were checked and met. In addition we used a nonparametric test, Kendall’s t rank correlation, to determine if relationships exist between biological and envi- ronmental variables and foxtail pine recruitment for northern and southern populations. All Statistical analyses were conducted with the software program JMP, version 8.0.1 (SAS Insti- tute Inc. Cary, NC). Current Population Trends Population trends were assessed by employing transition matrix models for each of the six populations of foxtail pine. In our study, transition matrices are used to describe and summarize current trends in survivorship, mor- tality, fecundity, and to a much lesser extent growth, as this is a long-lived tree species. Transition matrix models of populations follow the Lefkovitch (1965) model: n+) =An; where mn, is a column vector corresponding to the size structure at time t on the population classified into s size classes, and A is the matrix representing population dynamics. A is influ- enced by survival, growth, and reproduction. Entries in the transition matrix represent the 238 Size class 2 Size class 3 G; 5.1-10.0 P; P> P3 Fic. 2. MADRONO [Vol. 58 Size class 4 Size class 5 Size class 6 10.1-20.0 20.1-40.0 * 40.1 Py Ps Ps Size-class transition model for foxtail pine populations in California. Size classes are represented by circles with size class | at the left and proceeding to size class 6 at the right. Transition probabilities for growth (G;-G;) are the forward horizontal arrows, survivorship (P7-Ps5) are the bold circular arrows, and fecundity (F,-F,) are the long arrows from right to left. contribution each size class makes to every other size class during a specified time interval. We created sized-based models with six size classes: 1) recruits (individuals <1.37 m in height); 2) 0.1—5.0 d.b.h.; 3) 5.1-10 d.b.h.; 4) 10.1—20.0 d.b.h.; 5) 20.1-40.0 d.b.h.; and 6) =40.1 d.b.h. (Fig. 2). Transition probabilities were calculated and esti- mated from our field data. For growth probabil- ities, we assume that individuals will transition into the next size class if trees are in the upper d.b.h. limit of the size class, or height for seedlings/ saplings. For example, an individual with a d.b.h. of 4.9 cm or 5.0 cm would grow into the next size class in the next time step. Where mortality occurred, survivorship was calculated using the number of standing dead trees (years dead ranged from 1—12 yr) divided by current live and dead stems for each size class. In demographic studies of forest trees where mortality was not observed, mortality was assumed to be 0% or 2% (Ettl and Cottone 2002; van Mantgem et al. 2004). In our populations where no mortality was observed, we assumed a minimal value of 1% mortality; an average of these previously published estimates. Fecundity was estimated from existing recruitment and cone production data we collected for size classes 4, 5, and 6. We used the following formula to estimate fecundity for each of the 3 size classes: (no. of cones in stage(¢)/no. of cones for population(t)) x (no. of recruits for population(f)) no. of trees in stage(f) where ¢ is time. This formula is similar to that used by Davelos and Jarosz (2004) for estimating reproduction for American chestnut (Castanea dentata Marsh.). The population growth rate (A), as estimated using the dominant eigenvalue of the transition matrix (Caswell 2001), measures the rate of change in total population size. Population growth rate is a function of size- or age-specific rates of survival, growth and reproduction, with X > | indicating growing, A = | indicating stable, and A < 1 indicating declining populations. Classical statistical tests using estimates of A are inappropriate because demographic parameters and estimates of ~ are not simple and _ their distributions are often not known (Caswell 2001). Therefore, we computed 95% confidence inter- vals for X by bootstrapping (n = 10,000) across survivorship, growth, and fecundity estimates comprising the transition matrix. All calculations and bootstrapping were performed in Matlab (Mathworks Inc., Natick, MA). Kendall’s t rank correlation analysis was used to determine if relationships existed between biological and environmental variables and mean fecundity and survivorship. This nonparametric test was conducted with the software program JMP, version 8.0.1 (SAS Institute Inc. Cary, NC). RESULTS Forest and Stand Conditions Locations ranged in elevation from 2086 to 3398 m, with stands varying in physiognomy and environmental characteristics (Table 1). Slopes ranged from 11 to 22 percent and aspect varied as well (Table 1). Demographic sites in the moun- tains of interior northwestern California receive high amounts of rainfall, ranging from 1051 mm to 1388 mm, compared to the southern Sierra Nevada with a range from 356 mm to 721 mm (Table 1). January minimum and July maximum temperatures also differ between the two regions, with warmer temperatures and relatively longer growing seasons in the north compared to the southern Sierra Nevada (Table 1). Relative hu- midity is on average higher in the north than in the south (61% vs. 44%; see Table 1). The percent 2011] MALONEY: ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER — 239 TABLE 1. CLIMATE AND GEOLOGY SUMMARIES FOR FOXTAIL PINE POPULATIONS. Climate averages for each population based on PRISM data for 30-year averages from 1970-2000 (Daly et al. 1994 and FHTET). Ann. ppt = total annual precipitation in millimeters (mm), average minimum temperature for January and average maximum temperature for July in degrees Celsius, GDD growing-degree days, rel. humidity (%) = percent relative humidity, % max. rad. input (Cal. cm */year ) = percentage maximum solar radiation input was calculated using slope and aspect (Buffo et al. 1972). Soil data source: USDA NRCS (2008). Northern Southern North Yolla Onion Cottonwood _ Sirretta Lake Mountain Mount Eddy Bolly Valley Pass Peak Elevation (m) 2086 2453 2246 3073 3398 2925 Slope (%) 19 14 22 19 11 14 Aspect 133 135 208 27 207 250 Ann. ppt. (mm) 1313 1051 1388 356 439 121 Avg. min. temp. ("C) =—[e/ —3.1 =.) cael A) —9.6 — 9.) Avg. max. temp. (°C) 24 29 26 29 16.7 21 May GDD 48 0 32 410 0 0 September GDD 323 83 19] 131 0 145 Rel. humidity (%) 65 58 59 3] 45 57 % max. rad. input 85.39 92.36 95.89 54.42 90.61 85.02 (Cal. cm ?/year) Geology/parent gneiss/ serpentine/ metamorphic granitoid subglacial residuum material metamorphic/ metavolcanics rock till/granite | weathered serpentine granite maximum radiation input (Cal.cm */yr) was higher in the northern foxtail stands, mean = 91.20, than in the southern Sierra Nevada, mean = 78.02 (Table 1). The geology of the Klamath region is very diverse and sites in the north span a range of geological substrates (gneiss, serpentine, metamorphic, and metavolcanics) versus the southern Sierra Nevada sites that are primarily on granite (Table 1). White pine blister rust was only found in populations in the north, at Lake Mountain and Mount Eddy, with no rust in the Southern Sierra Nevada (Table 2). Mountain pine beetle was observed in 4 of the 6 sites with the highest incidence at Sirretta Peak followed by Lake Mountain (Table 2). Moderate mortality was found in 5 populations ranging from 0 to 14% TABLE 2. A = absence from stand. with an average of 5.7% in the north and 9% in the southern Sierra Nevada (Table 2). Clark’s nutcracker was observed in 1 of 3 stands in the north and present in all 3 popu- lations in the southern Sierra Nevada (Table 2). The only population in which Clark’s nutcracker was Observed in the north, at Mount Eddy, is also the only northern foxtail stand in which white- bark pine is an associate species. Whitebark pine is one of the preferred food resources for this corvid (Hutchins and Lanner 1982). Evidence of fire was found in 38% of the plots in the north and 88% in the southern Sierra Nevada (Table 2). In the southern Sierra Nevada, fire may be a relatively common disturbance agent in foxtail pine stands (Rourke 1988; North et al. 2009). BIOLOGICAL AND ENVIRONMENTAL SUMMARIES FOR FOXTAIL PINE POPULATIONS. P = presence and Northern Southern Lake Mount North Yolla Onion Cottonwood Sirretta Mountain Eddy Bolly Valley Pass Peak Pinus balfouriana density (inds. ha ') 89 95 64 114 224 46 Pinus balfouriana basal area (m° ha ') 10.6 26.4 10.8 25 43.2 13.4 Average Pinus balfouriana d.b.h. 249 45.2 35.3 44.1 359 50.9 Reproductive adults (inds. ha“ ') 30 49 28 62 138 35 Cones (no. ha ') 3966 3657 3463 To25 11,078 A325 Recruit. (inds.ha ') 64 370 74 40 227 7 Clark’s nutcracker A P A P P P WPBR (%) 4 2 0 0 0 0 MPB (%) 4 2 0 ] 0 16 Mortality (%) 13 4 0 6 7 14 Rock cover (%) 25.3 55.0 45.5 50.0 38.7 25.0 Evidence of fire (freq.) 33 50 33 100 66 100 240 MADRONO Composition and Structure Mean density of foxtail pine in northern stands was 82.6 trees/ha and ranged from 64 to 95 trees/ ha (Table 2). In southern foxtail pine stands the mean density was 129 trees/ha and ranged from 46 to 227 trees/ha (Table 2). Mean basal area was higher in the southern stands, 27.9 m’/ha, but ranged from 13.4 to 43.2 m’/ha. Mean basal area in northern foxtail pine stands was 15.9 m’/ha and ranged from 10.6 to 26.4 m’/ha (Table 2). Relatively higher basal areas in southern foxtail pine stands corresponded with a larger average d.b.h. of 43.6 cm (range: 35.9 to 50.9), compared to the north, 36.1 cm (range: 27.9 to 45.2; Table 2). Foxtail pine is the dominant component in these stands with importance values ranging from a 56.4% to 100% (Table 3). Some common associ- ates include red fir, western white pine, white- bark pine, limber pine and white fir (Table 3). Total tree density (all species) ranged from 67.3 to 227.0 individuals/ha and basal area ranged from 10.2 to a high of 43.2 m’/ha (Table 3). The size structure for northern populations is relatively mixed across diameter classes with the exception of Mount Eddy, which has higher numbers of trees in the largest size class (Fig. 3a). Size structure in the southern foxtail populations was generally skewed, with lower numbers in the smaller size classes and higher numbers in the largest size class (Fig. 3b). However, Cottonwood Pass has good representation of trees in the smallest size class (Fig. 3b). Reproductive Output and Recruitment Patterns Southern foxtail stands had higher numbers of reproductive individuals, mean = 78.3, than northern stands, mean = 35.7 (Table 2). Higher numbers of reproductive adults corresponds with higher cone production in the south, mean = 7642 cones/ha, and in the northern populations cone production averaged 3695 cones/ha (Table 2). However, what appears to be somewhat higher reproductive output in the south does not correspond to higher numbers of seedlings and saplings. Regeneration in the north averaged 169.3 seedlings/saplings per hectare compared to 91.3 seedlings/saplings per hectare in the southern populations (Table 2). Regional climate and landscape characteristics may strongly influence recruitment patterns, but microenvironmental conditions may be as influ- ential in the successful establishment of foxtail pine as large-scale phenomena. Foxtail seedlings and saplings in the north were generally growing on microsites with low litter depths, on rocky substrates, and in open canopies (Table 4). In the southern foxtail pine stands, recruitment was growing more frequently on microsites with higher amounts of litter, decomposed granite, in STAND STRUCTURE AND COMPOSITION FOR SIX PINUS BALFOURIANA POPULATIONS. Relative density = RD, relative basal area = RBA, and importance value = IV of tree species =1.37 m. Importance values were calculated for each species as (relative density + relative basal area)/2. TABLE 3. Southern Northern Mount Eddy Sirretta Peak Cottonwood Pass Onion Valley North Yolla Bolly Lake Mountain RD RBA IV% RD RBA IV% RD RBA IV% RD RBA IV% RD_ RBA IV % RD RBA IV % Species 0.3 223 >0.5 279 0.6 36.7 Abies concolor White fir Abies magnifica Red fir She, 13:5 16.1 214 us albicaulis Whitebark pine nN P [Vol. 58 W \O eT Ea oe: \O (oa) N —<+t Va) = CON S Vay — No) | | LV ay — =) (=) tC Klan co | WY = oN 2 i) | se = N S S| | || (=) ee Coo [~~ na oS | | jad = or \O ~~ xo | | | (oe) Saal \O “ll files sll || [on all | ~~ \O a on | nl|an ~~ CON Va) Via) Ss] |] uw [on oo & el | (=) A OMAN CO No) | Soot \O So fore) a = Fee N CO \O S| ea Se) te ov QY a c ha om! an ¢ 8 BoE AT s~aao . | S2nn 8S moses Rew S48. §4a 3 —_| = San Sets &.3¢ SSHE S25 SESS S58 SSeiyg sus Seeg8Saq EUROS ues leo 6 aad Aew 2011] MALONEY: ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER — 241 Said @ Lake Mountain Mt Eddy @ North Yolla Bolly 80.00 8 70.00 x —| 7 60.00 Q. < 50.00 = = 40.00 Same Fs 30.00 = ; | Zz 20.00 a 0.00 0.1-5.0 5.1-10.0 10.1-20.0 20.1-40.0 >40.1 A Diameter size class distribution (cm) HB Onion Valley Cottonwood Pass @ Sirretta Peak 90.00 » 80.00 = y 70.00 x — 2 60.00 2 : N E 50.00 - S 40.00 : fmt Ba s “e e = 30.00 4 : = ee Le a ‘“ 20.00 10.00 a a as oa 0.00 | 0.1-5.0 5.1-10.0 10.1-20.0 20.1-40.0 >40.1 B Diameter size class distribution (cm) FIG. 3. an open canopy, and associated with a rock shelter microhabitat (Table 4). No relationships were found between biologi- cal and environmental variables and foxtail pine recruitment from demographic plots in the northern region (see Table 5). In the north the Size structure of six foxtail pine populations from the northern (A) and southern (B) regions. best relationship, albeit weak, was between recruitment and foxtail pine density (Table 5). Density of foxtail pine may correspond with dominance in a stand and possibly an increase in the source of propagules (e.g., seed) as well as a higher competitive advantage. Two of the three 242 MADRONO TABLE 4. [Vol. 58 SUMMARY OF DOMINANT MICROENVIRONMENTAL CONDITIONS OF P. BALFOURIANA RECRUITMENT (INDIVIDUALS <1.37 MIN HEIGHT). Litter depth, in centimeters, surrounding seedling; substrate regeneration is growing in: es = exposed soil (no litter), dg = decomposed granite, sl = soil + litter, lg = log, rk = rock, or combinations dg] = decomposed granite + litter; canopy conditions: | = open/exposed, 2 = partially closed, 3 = closed; microhabitat conditions: 1 = tree/shrub nurse, 2 = rock shelter, 3 = other, 4 = none. Average recruitment ha“! 169.3 9153 Litter depth (cm) 0,52 2.40 North South study sites are located in areas with mixed geological sources including serpentine/ultra- maphic substrate (see Table 1). If foxtail pine is dominant at a site with serpentine soils it may have a competitive advantage in recruiting more successfully than other conifer associates, partic- ularly shade-intolerant species such as Abies concolor (Gordon & Glend.) Hildebr., A. magni- fica and Tsuga mertensiana (Eckert and Eckert 2007; see also Table 3). Strong positive relationships were found in the southern Sierra Nevada between foxtail recruit- ment and foxtail density, number of cones ha‘', and number of reproductive adults; all corre- sponding with source strength and propagule pressure (Table 5). A strong negative relationship was found between recruitment and tree species richness (Table 5). In the southern Sierra Nevada foxtail pine is often the dominant species, but in tree species-rich subalpine forests it may be at a competitive disadvantage in successfully recruit- ing into limited and favorable microsites (see Tables 3 and 4). Despite differences in factors potentially im- portant to recruitment of foxtail pine in the north and south, establishment patterns of foxtail pine in both regions appear to be somewhat episodic (Fig. 4). In the north, Mount Eddy appears to be recruiting relatively consistently since 1970, with a pulse of recruitment from 1998 to present (Fig. 4A). Recruitment number and patterns for both Lake Mountain and the North Yolla Bolly TABLE 5. Microenvironment Substrate (freq.) Canopy (freq.) Microhabitat (freq.) 64 (rk) 94.8 (open) 86.7 (none) 54 (dgl) 83.3 (open) 78.1 (rock shelter) are generally low and episodic (Fig. 4A) as is the case with the southern Sierra Nevada (Fig. 4B). Cottonwood Pass is the only population in the south that has had a recent pulse of recruitment from 1998 to present (Fig. 4B). Recruitment at Onion Valley is fairly low and episodic and recruitment is almost non-existent at Sirretta Peak, and any individual <1.37 m in height recruited prior to 1970 (Fig. 4B). Current Population Trends Foxtail pine populations varied in fecundity, survivorship, and growth (Table 6, Appendix 1). Fecundity was quite variable and this is reflected in the variation we found in cone production, number of reproductive adults, and the number of seedlings that successfully established in each of these populations (Tables 2 and 6). For both northern and southern foxtail pine populations, fecundity was positively correlated with solar radiation input and foxtail pine density (t = 0.466, P = 0.188; t = 0.466, P = 0.188, respectively). Survivorship varied, due to differ- ential mortality between populations (Tables 2 and 6), with trees in the largest diameter class (=40.1 cm dbh) having relatively lower survival rates. Low estimates of survivorship were found at Lake Mountain and Sirretta Peak (Table 6, Appendix 1). A negative correlation was found between foxtail survivorship and incidence of MPB (t = —0.41, P = 0.251). Growth 1s the least KENDALL’S t RANK CORRELATIONS BETWEEN BIOLOGICAL AND ENVIRONMENTAL VARIABLES AND FOXTAIL PINE RECRUITMENT FOR NORTHERN (N = 8 PLOTS) AND SOUTHERN POPULATIONS (N = 8 PLOTS). Northern Southern Variable t P-value 5 P-value Ann. ppt. 0.000 1.000 0.000 1.000 Max July temp. 0.178 0.584 —(0.481 Olea No. cones =0.231 0.441 0.691 0.017 Solar radiation —0.077 0.797 0.109 0.708 Tree species richness 0.136 0.675 —0.645 0.041 Pinus balfouriana density 0.353 0.244 0.764 0.008 Pinus balfouriana basal area —0.077 0.797 0.545 0.061 No. reproductive adults —0.196 0517 0.618 0.034 Litter depth —0.105 0.800 0.195 0.534 Rock cover 0.117 0.698 —(). 109 0.708 2011] MALONEY: ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER — 243 10 OC) Lake Mtn @ Mt Eddy O North Yolla Bolly 2002 2003 2004 2005 2006 2007 2008 2000 2001 10 CJ Onion Valley @ Cottonwood Pass(C Siretta Peak 2003 Ln] 2008 L) 2000 2001 2002 2004 2005 2006 2007 Year Fic. 4. Foxtail pine recruitment and establishment patterns for populations in the north (A) and south (B). dynamic and variable transition element, given 1.0, indicating that most of these populations the long-lived nature of this tree species and a appear stable (Table 6). The North Yolla Bolly one-time sampling (Table 6, Appendix 1). population had the highest A, followed by Onion Estimates of population growth rate (A) varied Valley, Mount Eddy, Cottonwood Pass, and among foxtail sites, with 5 of the 6 havinga’= __ Sirretta Peak (Table 6). Noteworthy of the North 244 TABLE 6. MADRONO [Vol. 58 MEANS FOR ESTIMATED FECUNDITY, SURVIVORSHIP, AND GROWTH FROM SIZE-BASED TRANSITION MATRICES FOR SIX FOXTAIL PINE POPULATIONS. Estimated population growth rate, A, is shown along with 2.5% and 97.5% confidence intervals (CI, in parentheses). Population Fecundity Survival Lake Mountain 0.233 0.898 Mount Eddy 0.677 0.971 North Yolla Bolly 0.208 0.976 Onion Valley 0.082 0.966 Cottonwood Pass 0.242 0.950 Sirretta Peak 0.026 0.906 Yolla Bolly population is that it 1s the second smallest population with 141 individuals/ha (Sirretta Peak was the smallest with 54 individ- uals/ha), but it is the one population where no WPBR, MPB, or mortality was observed, with consistent and high survivorship across all size classes (Table 6, Appendix 1). Lake Mountain has a A < 1.0, indicative of a population that may be in decline. At Lake Mountain both WPBR and MPB were present, with lower survivorship rates in size classes 3, 5, and 6 (Table 6, Appendix 1). All upper limit confidence intervals for A were greater than 1, suggesting stability. However, for most populations the lower confidence interval was <1.0 (Table 6); whether this is cause for concern is difficult to assess, given that > was estimated from a one-time sampling. DISCUSSION Ecological and environmental conditions clear- ly differ between the two regions in which foxtail pine grows, influencing species, stand, and demographic characteristics. In the north, popu- lations exist under a more mesic climatic regime, a diversity of geological substrates, stands of high tree species richness, lower densities and basal areas, but higher numbers of recruitment, with a relatively mixed size class distribution. Southern foxtail pine populations exist under a more xeric climatic regime dominated by granitic substrates, with moderate to high stand densities, less tree species-rich, lower recruitment numbers, but higher basal areas; largely due to a skewed size class distribution with high representation of large diameter trees. Fecundity, a key demographic parameter in forest tree populations (see Clark et al. 1999), in this study was a function of the number of cones produced in a population and the number of recruits successfully established. At the stand or population-level, solar radiation input and foxtail pine density were correlated with fecundity across all locations, north and south. These factors may correspond with microenvironmental and topo- graphic conditions that favor germination (e.g., warmer microclimate) in subalpine environments as well as propagule pressure (e.g., seed supply), which increases with tree size (basal area). Large- Growth d (2.5%, 97.5% CI) 0.026 0.998 (0.588, 1.576) 0.012 1.013 (0.677, 1.552) 0.053 1.041 (0.703, 1.527) 0.040 1.019 (0.775, 1.497) 0.025 1.011 (0.419, 1.720) 0.066 1.007 (0.705, 1.571) scale phenomena such as regional climate and landscape features are known to influence re- cruitment in subalpine forests (Millar et al. 2004, 2006; Bunn et al. 2005). But at the plot-level, microenvironmental conditions (e.g., litter, sub- strate type, and microhabitat) and factors corre- sponding to local seed supply (e.g., density, basal area, number of cones, and number of reproduc- tive adults) may be equally as important to successful recruitment of foxtail pine. An important limitation of this study is a lack of information about seed dispersal or seed/cone predation, both important factors in recruitment dynamics. Wind is the primary dispersal mecha- nism of foxtail pine seed (see Mastrogiuseppe and Mastrogiuseppe 1980). However, the presence of Clark’s nutcracker in all the southern Sierra Nevada populations, even in the absence of whitebark pine, raises the question of the role of bird and/or small mammal dispersal. Clark’s nutcrackers were observed in one of three sites in the Klamath region but this bird has been observed feeding on foxtail pine seed in this region, in sites where whitebark pine is not present (A. Eckert, Virginia Commonwealth Univ., personal communication). The closely- related Great Basin bristlecone pine (P. /ongaeva) resides in nearby mountain ranges (White, Inyo, and Panamint) to the east of foxtail pine in the southern Sierra Nevada. While Great Basin bristlecone pine is mainly dispersed by wind, Lanner (1988) and Lanner et al. (1984) have shown that Clark’s nutcrackers can also play a role in dispersal and subsequent regeneration of this enigmatic high-elevation white pine. Seventy- eight percent of foxtail pine recruits in the southern Sierra Nevada were growing in a rock shelter (Table 4). While soil moisture availability and shade provided by a rock shelter can favor seed germination and seedling survival, Clark’s nutcrackers are also known to select cache sites next to rocks (Tomback 1978). Another impor- tant factor influencing recruitment may be insect predation of cones and seed, which has been largely overlooked, but may have an overwhelm- ing effect on cone production and seed supply in some years. In 2009 and 2010 a high frequency of insects (e.g., Dioryctria spp.) were observed in cones of foxtail pine in the southern Sierra 2011] MALONEY: ECOLOGY AND DEMOGRAPHY OF AN ENDEMIC SUBALPINE CONIFER Nevada and Klamath region (D. Davis, A. Delfino Mix, P. Maloney, and D. Welty, personal observations). Both dispersal dynamics and cone/ seed predation are areas in recruitment studies of forest trees that warrant further investigations. Recruitment patterns of foxtail pine appear to be episodic in nature and other studies have observed this temporal trend for foxtail pine and other subalpine conifers (Millar et al. 2004, 2006; Bunn et al. 2005). Unfortunately this study lacks yearly temperature and precipitation data for these two regions that might be associated with recruitment years. Warmer temperatures and above average precipitation years can influence recruitment pulses (Millar et al. 2004, 2006; Bunn et al. 2005) and possibly the episodic recruitment patterns observed in this study. Foxtail pine is recruiting in relatively higher numbers in the north than in the southern Sierra Nevada. But despite lower numbers of observed recruitment in some locations, most populations (north and south) appear to be buffered from declines due to high survivorship across all size classes, including larger reproductive individuals. Four of the six populations, two in the north and two in the south, appear to be stable, due to low mortality and high survival. Lake Mountain (A = 0.998) and Sirretta Peak (A = 1.007) have low estimated growth rates, due to low survivorship of individuals in the largest size class (0.786 and 0.762, respectively). The incidence of MPB was much higher and more frequently observed in the southern Sierra Nevada than in northern stands. Drought conditions appear to trigger MPB activity in the high elevation white pine forests of California (CFPC Reports 1976-2009; Millar et al. 2007). Because little is known about MPB in high elevation forests of California or about historical outbreaks, it is difficult to say what might be out of the range of historical variability for this native insect. Certainly old dead snags are observed in these forests but the causes of death are unknown. MPB-mediated mortality coupled with drought stress may be important factors contributing to current population trends at Lake Mountain and Sirretta Peak. Interestingly each of these populations represents the northern range limit (Lake Mountain) and southern range limit (Sirretta Peak) for the two subspecies in north- western California and the southern Sierra Nevada, respectively (see Fig. 1). The presence of WPBR at Lake Mountain may also be a predisposing factor influencing survivor- ship. In California, WPBR has only been found in northern foxtail pine stands and has not been confirmed in the southern stands of foxtail pine (Maloney, 2011). Latitudinal trends in WPBR incidence may correspond with a longer residence time for C. ribicola in the north (1929-1938) compared to the southern Sierra Nevada (1961) (Smith 1996). Climatic conditions in northern 245 stands of foxtail pine are more favorable for C. ribicola infection (e.g., higher annual rainfall, warmer temperatures, and higher relative humid- ity), whereas the environment of the southern subspecies is drier and colder. We know that WPBR is present lower in elevation in Sequoia and Kings Canyon NP and Sequoia NF, but given the environment of the high Sierra Nevada where foxtail pine grows, these conditions may limit C. ribicola spread into these high elevation forests (see Maloney 2011). Because population growth, A, was calculated from a One-time sampling, our estimates may not reflect intrinsic variation in rates of fecundity, survivorship, mortality, and growth. Year to year variation in climate, cone production, cone and seed predation, recruitment success, tree mortal- ity, insect dynamics, fire activity, and conditions favorable for WPBR infection can be consider- able. Our intent was not to predict future popu- lation growth but to describe current population conditions and trends. Another limitation of our study is that the confidence intervals are very large for A, which is a cause for concern, but it is difficult to assess the magnitude of this concern as was estimated from a one-time sampling that likely resulted in the large variances around the point estimate. Obtaining long-term demographic data for long-lived tree species is difficult, but will be critical to accurately access population dynamics of foxtail pine in an era of rapidly changing climate and increasing environmental stressors (e.g., insect outbreaks and non-native diseases). Ecological and environmental differ- ences between the northern and southern subspe- cies may influence how P. balfouriana responds to natural and anthropogenic disturbances. Such information is basic to developing conservation, monitoring, and management strategies for this endemic and narrowly distributed white pine in subalpine forests of California. ACKNOWLEDGMENTS The author would like to thank USDA Forest Service cooperators: Joan Dunlap, Lisa Fischer, Deems Bur- ton, Phil Cannon, Hugh Safford, and Michael Bohne as well as National Forest Wilderness, District Rangers, and Forest botanists; especially Jeff Novak and Kathleen Nelson of the Inyo NF. The author thanks Camille Jensen, Shannon Lynch, Allison Wickland, Tom Burt, and Hugh Denno for field support and Akiko Oguchi for GIS maps. A special thank you to Andrew Eckert, Deems Burton, and Dean Davis for sharing their knowledge of Pinus balfouriana ecology, genetics, and conservation. We also thank the following people and agencies for providing data used in this project: USDA FS, Forest Health Technology Enter- prise Team (FHTET), Fort Collins, CO, F. Krist and L. Lewis; USDA Natural Resources Conservation Service (NRCS), South Lake Tahoe CA, W. Loftis. The author appreciates the comments made on an earlier draft by Marc Meyers, Hugh Safford and Phil Cannon. 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Effects of an introduced pathogen and fire exclusion on the demography of sugar pine. Ecological Applications 14:1590—1602. 248 MADRONO [Vol. 58 APPENDIX 1 Size-based transition matrices for recruitment (<1.37 meters in height) and stem diameters at breast height (1.37 m) in centimeters for six foxtail pine populations in California. Transition probabilities for survivorship and growth are in bold and shaded cells, respectively. Fecundity estimates are in italics. Size year n Size description Sizen+t 1 5) 2 4 5 6 Lake Mountain, n = 153 recruitment 1 0.992 0 0 0.018 0.206 0.475 0.1—5.0 dbh 2 0.008 0.994 0 0 0 0 5.1-10.0 dbh 3 0 0.006 0.833 0 0 0 10.1—20.0 dbh 4 0 0 0.056 0.947 0 0 20.140 dbh 5 0 0 0 0.053 0.833 0 =40.1 dbh 6 0 0 0 0 0.006 0.786 Mount Eddy, n = 466 recruitment l 0.999 0 0 0.034 0.658 1.338 0.1-5.0 dbh 2 0.027 0.995 0 0 0 0 5.1-10.0 dbh 3 0 0.005 0.993 0 0 0 10.1—20.0 dbh 4 0 0 0.007 0.988 0 0 20.140 dbh 5 0 0 0 0.013 0.917 0 =40.1 dbh 6 0 0 0 0 0.008 0.934 North Yolla Bolly, n = 141 recruitment l 0.990 0 0.056 0.111 0.458 0.1—5.0 dbh 2 0.010 0.989 0 0 0 0 5.1-10.0 dbh 3 0 0.100 0.900 0 0 0 10.1—20.0 dbh 4 0 0 0.100 0.988 0 0 20.1—40 dbh 5 0 0 0 0.013 0.996 0 =40.1 dbh 6 0 0 0 0 0.042 0.994 Onion Valley, n = 154 recruitment ] 0.988 0 0 0.071 0.079 0.096 0.1-5.0 dbh 2 0.013 0.990 0 0 0 0) 5.1-10.0 dbh 3 0 0.100 0.986 0 0 0 10.1—20.0 dbh 4 0 0 0.014 0.909 0 0 20.1—40 dbh =) 0 0 0 0.045 0.972 0 =40.1 dbh 6 0 0 0 0 0.028 0.952 Cottonwood Pass, n = 453 recruitment l 0.998 0 0 0.063 0.218 0.445 0.15.0 dbh 2 0.002 0.998 0 0 0 0 5.1-10.0 dbh 3 0 0.020 0.995 0 0 0 10.1—20.0 dbh 4 0 0 0.045 0.903 0 0) 20.140 dbh 5 0 0 0 0.032 0.905 0 =40.1 dbh 6 0 0 0 0 0.024 0.902 Sirretta Peak, n = 54 recruitment ] 0.900 0 0 0.008 0.024 0.046 0.1—5.0 dbh 2 0.100 0.900 0 0 0 0 5.1-10.0 dbh 3 0 0.100 0.900 0 0 0 10.1—20.0 dbh 4 0 0 0.100 0.983 0 0 20.140 dbh 5 0 0 0 0.017 0.988 0 =40.1 dbh 6 0 0 0 0 0.013 0.762 MADRONO, Vol. 58, No. 4, pp. 249-255, 2011 MOLECULAR PHYLOGENETICS OF GARRYA (GARRYACEAE) DYLAN O. BURGE! Duke University Department of Biology, Box 90338, Durham, NC 27708 dylan.o.burge@gmail.com ABSTRACT Garrya (Garryaceae) comprises 15 species of shrubs and small trees restricted to the Americas. Garrya is taxonomically divided into two subgenera, Garrya and Fadyenia, which differ in morphology, secondary chemistry, and geographic distribution. The present work uses nuclear ribosomal DNA (ITS) sequence data from 11 Garrya species to elucidate phylogenetic relationships within the genus and test the monophyly of the subgenera. Results strongly support subgenus Fadyenia as monophyletic, while monophyly of subgenus Garrya is supported only by maximum parsimony analyses. ITS data do not provide evidence for genetic admixture between the two subgenera of Garrya in spite of broad geographic overlap. Key Words: Aucuba, California, Eucommia, Fadyenia, Garrya, Garryaceae, ITS, Mexico. Garrya Douglas ex Lindl. contains 15 species endemic to North America, Central America, and the Caribbean (Dahling 1978; Nesom unpub- lished; Table 1; Fig. 1). Garrya species are dioecious shrubs and small trees with decussate, evergreen leaves and pendulous, catkin-like inflorescences. The plants are probably wind pollinated (Hallock 1930; Dahling 1978; Liston 2003). Garrya is found in a diversity of habitats over its broad geographic range, from cloud forest to maritime chaparral, but is typically a component of shrublands (e.g., chaparral) or forests (Dahling 1978). Molecular phylogenetic studies consistently resolve Garrya as sister to the east Asian shrub genus Aucuba (Soltis et al. 2000; Bremer et al. 2002), which together comprise Garryaceae (APG 2003). These results confirm a close relationship that has long been hypothesized on the basis of morphology and chemistry (reviewed in Liston 2003). Molecular phylogenetic studies also support a close relationship between Gar- ryaceae and the monotypic east Asian tree Eucommia (Eucommiaceae), which together com- prise the euasterid order Garryales (APG 2003). Garrya 1s divided into two subgenera, Garrya (6 spp.) and Fadyenia (9 spp.), which differ in geographic distribution, inflorescence morpholo- gy, and secondary chemistry (Dahling 1978; Table 2). Subgenus Fadyenia has its center of diversity in Mexico, while subgenus Garrya reaches peak diversity in the western U.S. The geographic distribution of the subgenera overlaps in the southwestern U.S. and Mexico (Dahling 1978). Research presented here aims to elucidate phylogenetic relationships in Garrya and relate 'Present address: National Herbarium of New South Wales, Royal Botanic Gardens, Sydney, Mrs Mac- quaries Road, New South Wales, 2011, Australia. this to the taxonomy, distribution, and biology of the species. Specifically, I test the hypothesis that the Garrya subgenera are monophyletic, with separate histories of diversification in different geographic regions of the Americas. MATERIALS AND METHODS Genetic Sampling Sampling of Garrya populations was designed to represent the geographic range of the genus with an emphasis on California, the southwestern U.S., and Mexico (Table 3; Appendix 1; Fig. 1). One sample of Aucuba was obtained from a garden planting in California. DNA from 22 Garrya individuals was studied, representing 11 of the 15 species currently recognized (Dahling 1978; Nesom unpublished). For the species occurring in the U.S., voucher specimens were identified according to Nesom (unpublished); for all exclusively Latin American taxa, identifica- tions were according to Dahling (1978). Voucher specimens are deposited at DUKE (Appendix 1). Molecular Methods Genomic DNA was extracted from silica-dried leaf tissue using the DNeasy Plant Mini Kit (Qiagen, Germantown, MD) according to the manufacturer’s instructions. Polymerase chain reactions were performed using Qiagen Taq DNA Polymerase. Amplification was performed using an initial incubation at 94°C for 10 min and 30 cycles of three-step PCR (1 min at 94°C, 30 sec at 55°C, and 2 min at 72°C), followed by final extension at 72°C for 7 min. I amplified the nuclear ribosomal ITS region (ITS 1, 5.8S, and ITS2) using the primers ITS4 (White et al. 1990) and ITSA (Blattner 1999). I amplified the trnL-F plastid region, comprising the ¢rnL intron and the 250 TABLE |. MADRONO [Vol. 58 GARRYA SPECIES AND SAMPLING. Sampling: number of populations sampled for phylogenetic analysis (Table 3); Distribution: geographic distribution of the species (SW U.S.: southwestern United States); CFP: indicates whether species occurs in the California Floristic Province; Flower: range of known flowering times for species (Dahling 1978). ° subspecies are recognized by Dahling (1978) and/or Nesom (unpublished): G. ovata: 3 subspecies; G. laurifolia: 4. Species Sampling Distribution CFP Flower Subgenus Fadyenia G. fadyena Hook. 0 Greater Antilles Dec-Feb G. glaberrima Wangerin 3 E Mexico Mar-May G. grisea Wiggins ] Mexico (Baja California) xX Feb-Apr G. laurifolia Benth.” 2 Mexico and Central America Dec-Apr G. lindheimeri Torr. l SW U:S., Mexico Mar-May G. longifolia Rose 0 Mexico Jan-Mar G. ovata Benth.® l SW U.S. and Mexico Mar-Apr G. salicifolia Eastw. 0 Mexico (Baja California Sur) Aug-Dec G. wrightii Torr. 3 SW U.S. and Mexico Apr-Aug Subgenus Garrya G. buxifolia A. Gray | U.S. (CA and OR) xX Feb-Apr G. corvorum Standl. & Steyerm. 0 Guatemala Dec-Jan G. elliptica Douglas ex Lindl. 2 U.S. (CA and OR) x Dec-Feb G. flavescens S. Watson 3 U.S. (AZ, CA, NV NM, UT) and »,« Feb-Apr Mexico G. fremontii Torr. 3 U.S. (CA, OR, WA) xX Jan-Apr G. veatchii Kellogg Zz U.S. (CA) and Mexico (Baja California) xX Jan-May trnL-F intergenic spacer, using primers c and f of Taberlet et al. (1991). Excess primer and dNTPs were removed using exonuclease I (New England Biolabs, Ipswich, MA [NEB]; 0.2 units/ul PCR product) and antarctic phosphatase (NEB; 1.0 unit/ul PCR product) incubated for 15 min at 37°C followed by 15 min at 80°C. For sequenc- ing, Big Dye chemistry (Applied Biosystems, Foster City, CA) was utilized according to the manufacturer’s instructions. Sequences were de- termined on an Applied Biosystems 3100 Genetic Analyzer at the Duke University Institute for Genome Science and Policy Sequencing and Genetic Analysis Facility. Sequences and Alignment A total of 23 ITS and 12 trnL-F sequences were generated for the present study. A preliminary alignment of trnL-F revealed that the nine sequenced Garrya species shared a nearly identi- cal sequence; there was just a single nucleotide substitution difference among the species, a change unique to G. grisea Wiggins (D.O. Burge 778; Table 3). In addition, two insertion/deletion events were present within Garrya, 1) a 1 bp length difference within the trnL intron, where a poly-T region was one bp longer in two accessions of G. elliptica Douglas ex Lindl. (D.O. Burge 382 & 386; Table 3) than in remaining Garrya, and 2) a 2 bp length difference in the ¢rnL-F intergenic spacer, where a poly-T region was two bp longer in members of subgenus Fadyenia relative to members of subgenus Garrya. Because of this low level of variation, trnL-F was not sequenced in additional plants, and was abandoned in favor of ITS for subsequent alignment and tree building. The 23 new ITS sequences (22 Garrya and 1 Aucuba) were supplemented with an ITS se- quence for Eucommia ulmoides Oliv. from Gen- Bank (Table 3). All DNA sequences were assem- bled and edited using Sequencher 4.1 (Gene Codes Corporation). Edited sequences were deposited in GenBank (trnL-F: JN234721-32; ITS: Table 3). ITS sequences were aligned using MUSCLE (Edgar 2004) under default settings. Due to ambiguity, a 22 bp region of ITSI was excluded from all subsequent analyses. The alignment was deposited in TreeBase (Study 11755). Phylogenetic Analysis Trees were reconstructed using Bayesian, maximum likelihood (ML), and maximum parsi- mony (MP) techniques. Trees were rooted using Eucommia ulmoides (APG 2003). Bayesian phy- logenetic analyses were conducted using the best- fit model of evolution from AIC output of the program MrModeltest (GTR + G; Nylander 2004). Sampling of trees was performed using the program MrBayes 3.0 (Ronquist and Huel- senbeck 2003). Three separate runs of 1 x 10° MCMC generations were performed using one heated and three cold chains, sampling every 1000 generations. Independent chains were in- spected for convergence (standard deviation of 2011] Legend O Collections €> Garrya Distribution — 200 km BiG. 1, BURGE: PHYLOGENETICS OF GARRYA Garrya distribution and sampling. Distribution of Garrya indicated by dark gray shading (data from participants of the Consortium of California Herbaria, 2011). Sampling locations indicated by white circles (Table 3). split frequencies nearing 0.001). Log-likelihood for the sampled tree was plotted and examined in Microsoft Excel to assess convergence and determine an appropriate burn-in period (Ron- quist and Huelsenbeck 2003). A total of 1 =< 10° generations (100 trees) were eliminated as burn- in, leaving 9 X 10° generations (950 trees) of explored tree space for computing branch TABLE 2. Character Q inflorescence Ovary appendage Paired floral bracts Flowers per pair Fusion of pair Size in Q Geographic distribution Small, epigynous Three Subgenus Garrya Compact, pendulous, unbranched Basally connate, forming a cup Reduced in size, not leaf-like Western U.S. and northern Mexico lengths and posterior probabilities (PP) of clades. Consensus phylograms were built for each of the three independent runs using MrBayes (Ronquist and Huelsenbeck 2003). Following inspection to verify similarity of the results, trees from all three runs were combined in a consensus phylogram. Maximum likelihood tree building was performed in GARLI v 1.0 MORPHOLOGICAL AND DISTRIBUTIONAL COMPARISON BETWEEN THE TWO SUBGENERA OF GARRYA. Description Subgenus Fadyenia Loose, erect, branched Large, foliaceous, partially adnate One Distinct to partially adnate basally At least proximal large and leaf-like Western U.S., Mexico, Central America, and Caribbean 252 TABLE 3. MADRONO [Vol. 58 COLLECTION NUMBER AND PROVENANCE FOR VOUCHER SPECIMEN (APPENDIX 1) AND GENBANK ACCESSION NUMBERS FOR ITS SEQUENCES. All vouchers deposited at DUKE. Taxon Collection number and provenance GenBank ITS Aucuba japonica Thunb. D.O. Burge 363, Butte Co., CA JN234733 Eucommia ulmoides A. Gray -- AY 650006 Garrya, subgenus Fadyenia G. glaberrima Wangerin D.O. Burge 1025, Hidalgo, Mexico JN234743 D.O. Burge 1216, Nuevo Leon, Mexico JN234744 D.O. Burge 1225, Tamaulipas, Mexico JN234745 G. grisea Wiggins D.O. Burge 778, Baja California, Mexico JN234746 G. laurifolia Benth. D.O. Burge 1218, Nuevo Leon, Mexico JN234747 D.O. Burge 1252, Chihuahua, Mexico JN234748 G. lindheimeri Torr. D.O. Burge 750, Travis Co., TX JN234749 G. ovata Benth. D.O. Burge 1221, Nuevo Leon, Mexico JN234750 G. wrightii Torr. D.O. Burge 934, Pima Co., AZ JN234753 D.O. Burge 1239, Durango, Mexico JN234754 D.O. Burge 1253, Chihuahua, Mexico JN234755 Garrya, subgenus Garrya G. buxifolia A. Gray D.O. Burge 1160, Josephine Co., OR JN234734 G. elliptica Douglas ex Lindl. D.O. Burge 382, Monterey Co., CA JN234735 D.O. Burge 386, Marin Co., CA JN234736 G. flavescens S. Watson D.O. Burge 370, Kern Co., CA JN234737 D.O. Burge 419, Yavapai Co., AZ JN234738 D.O. Burge 1036, Baja California, Mexico JN234739 G. fremontii Torr. D.O. Burge 353, Butte Co., CA JN234740 D.O. Burge 362, Humboldt Co., CA JN234741 D.O. Burge 1148, Tuolumne Co., CA JN234742 G. veatchii Kellogg D.O. Burge 378, San Luis Obispo Co., CA JN234751 D.O. Burge 1041, Baja California, Mexico JN234752 (Zwickl 2006). Two search replicates of 1 x 10° generations were performed in a single execu- tion with a random starting tree. Other param- eters were kept at default values. Statistical support was inferred with 100 replicates of bootstrap reweighting (Felsenstein 1985) using 5 x 10° generations per replicate. The majority rule consensus tree was calculated using the 100 best bootstrap trees. Maximum parsimony phylogenetic analysis was carried out using PAUP* v 4.0 (Swofford 2000). Heuristic search- es used 1000 random sequence addition repli- cates and tree bisection-reconnection branch swapping. Nonparametric bootstrap analysis (Felsenstein 1985) was conducted using 100 pseudoreplicates and heuristic settings with 10 random sequence addition replicates. In all MP analyses, gaps introduced by the alignment process were treated as missing data. RESULTS DNA Sequences The ITS region for A. japonica Thunb. was 605 bp in length. In Garrya this region varied from 622 to 624 bp; in all members of subgenus Garrya the region was 624 bp long while in subgenus Fadyenia it varied from 622 (G. grisea, D.O. Burge 778; Table 3) to 623 bp. The ITS alignment (TreeBase Study 11755) contained 696 characters, 22 of which were excluded (see above). Of the 674 included characters, 189 were variable and 48 were parsimony informative. Phylogeny Bayesian, ML, and MP analyses provided similar topologies and levels of support (Fig. 2; TreeBase Study 11755). Maximum parsimony analysis resulted in 147 equally parsimonious trees (length = 217, CI = 0.97, RI = 0.97). A total of eight nodes are found in the strict consensus of these trees (Fig. 2). Overall, Garrya is strongly monophyletic (Bayesian PP 0.99; MP bootstrap 100%; ML boostrap 95%; Fig. 2); subgenus Fadyenia is also strongly supported as monophyletic (Bayesian PP 1.0; MP bootstrap 100%; ML boostrap 100%), with several moder- ately-supported groupings within it. Though subgenus Garrya is monophyletic in the strict consensus cladogram from MP analysis, and receives 89% MP bootstrap support, this group is not supported in Bayesian or ML analyses. In addition, a grouping of Garrya ovata Benth. with Garrya lindheimeri Torr. is strongly supported (Bayesian PP 1.0; MP bootstrap 86%; ML bootstrap 84%), as in a clade containing all sampled populations of Garrya glaberrima Wan- gerin and one Garrya /aurifolia Benth. (Bayesian PP 0.99; MP bootstrap 93%; ML _ bootstrap 90%). However, none of the seven Garrya species 2011] <50/58 0.92 55/67 0.93 86/84 100/100 100 1.00 62/61 93/90 0.87 0.99 100/95 0.99 -89/<50__ <0.50 FIG. 2. BURGE: PHYLOGENETICS OF GARRYA @ G. laurifolia (1252) wnghtii (1239) wnightii (1253) grisea (778) wrightii (934) lindheimeri (750) ovata (1221) glaberrima (1225) laurifolia (1218) ejuexpe+ snuebqns glaberrima (1216) glaberrima (1025) veatchii (1041) veatchii (378) O 6 eliiptica (382) O 4. fremontii (353) O G eliptica (386) O G. flavescens (1036) O G. buxifolia (1160) O 4. fremontii (1148) O G. flavescens (370) e G. flavescens (419) O 4 fremontii (362) E. ulmoides 999 9 O99 O99 9 OD ®D ® 008000808080 808080 efues snuebans A. japonica Strict consensus of 147 equally parsimonious trees recovered in maximum parsimony (MP) phylogenetic analysis, with support values from 100 MP bootstrap replicates above branches, at left. Tree is rooted using E. ulmoides. Support from maximum likelihood bootstrap (above branches, at right) and Bayesian analysis (below branches) mapped on tree. Species names followed by D.O. Burge collection numbers. Open circles indicate collections obtained from within the California Floristic Province (CFP); dark circles are from outside of the CFP. represented by more than one sampled plant are recovered as monophyletic (Fig. 2). DISCUSSION Phylogenetic Relationships ITS trees strongly support subgenus Fadyenia, as circumscribed by Dahling (1978; Fig. 2). The Phylogenetic isolation of subgenus Fadyenia is supported by morphology, secondary chemistry (Dahling 1978), and geographic distribution (Table 2). By contrast, the monophyly of subge- nus Garrya is strongly supported only by maximum parsimony trees (Fig. 2). The lack of support for subgenus Garrya that is seen in maximum likelihood and Bayesian analyses may represent an artifact of analysis due to the small size of the ITS dataset. Future studies should utilize additional genes from both the chloroplast and nuclear genomes. The strong relationship between G. ovata and G. lindheimeri indicated by ITS is supported by the similar morphology of these species (Nesom - unpublished). Indeed, the similarities are so great that G. lindheimeri was treated as part of G. _ ovata, at the subspecies rank, by Dahling (1978). _ Nevertheless, the species are ecologically distinct _ over most of their geographic range, and remain morphologically distinct in the parts of northern Mexico where they occur sympatrically, though hybrids may occasionally form (Nesom unpub- lished). It is also noteworthy that no individual Garrya species is monophyletic (Fig. 2). One potential exception is G. glaberrima; the three included individuals of this species group together rela- tively strongly with a single individual of G. laurifolia (Fig. 2). The strong divergence of G. glaberrima from remaining members of subgenus Fadyenia is supported by the unusual morphol- ogy and phytochemistry of the species (Dahling 1978); presence of one individual of G. /aurifolia (D.O. Burge 1281) in this group might be explained by geneflow, as this individual was collected in an area where G. glaberrima occurs (D.O. Burge 1216; Table 3; Figs. 1 and 2). The overall lack of monophyly for individual species of Garrya is noteworthy as it is consistent with the action of incomplete lineage sorting (Maddi- son and Knowles 2006) due to shallow genetic divergence among species, possibly exacerbated by geneflow. Hybrids are not frequently observed in the wild (Dahling 1978; D.O. Burge, personal observation; but see Munz and Keck 1968), and the extent of geneflow among species of Garrya has never been directly studied. Thus, incomplete lineage sorting stands as the most probable 254 explanation for the general lack of phylogenetic cohesion among populations of individual spe- cies. Nevertheless, the present study does not include all species, and is based on a small sample of populations; analysis of additional species and populations might reveal greater phylogenetic affinity among populations of individual species. In addition, the present study is based on a very small sample of DNA sequence data; additional data, ideally from both the chloroplast and nuclear genomes, might provide greater phyloge- netic support for individual species. Diversification of Garrya in the Americas Subgenus Fadyenia represents a lineage that has diversified in the mountainous regions of the southwestern U.S., Mexico, Central America, and the Greater Antilles (Fig. 1). If subgenus Garrya is monophyletic, as suggested by some phyloge- netic analyses (Fig. 2), the group would represent a diversification that is focused in the California Floristic Province (CFP) of western North America (Table 1, Fig. 2). In spite of the wide geographic overlap of these two groups in the southwestern U.S. and Mexico, which should present opportunities for interbreeding, molecular phylogenetic results do not provide evidence for geneflow between the two subgenera of Garrya. It is possible that this lack of geneflow is driven by differences in flowering time, as indicated by a slight tendency toward earlier flowering in subge- nus Garrya as compared to subgenus Fadyenia (Table 1). This idea is supported by the observa- tion of staggered flowering time at several locations in the southwestern U.S. where mem- bers of each subgenus occur as part of the same plant communities (D.O. Burge, personal obser- vation). ACKNOWLEDGMENTS The author thanks the Duke University Systematics Discussion Group for constructive criticism of this work. The author also thanks Katherine Zhukovsky and Kaila Mugford for assistance with lab work. Funding was provided by the Duke University Depart- ment of Biology. LITERATURE CITED ANGIOSPERM PHYLOGENY GROUP, THE (APG). 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141:399-436. BLATTNER, F. R. 1999. Direct amplification of the entire ITS region from poorly preserved plant MADRONO [Vol. 58 material using recombinant PCR. Biotechniques 27:1180-1185. BREMER, B., K. BREMER, N. HEIDARI, P. ERIXSON, R. G. OLMSTEAD, A. A. ANDERBERG, M. KAL- LERSJO, AND E. BARKHORDARIAN. 2002. Phyloge- netics of asterids based on 3 coding and 3 non-coding chloroplast DNA markers and the utility of non- coding DNA at higher taxonomic levels. Molecular Phylogenetics and Evolution 24:274-301. DAHLING, G. V. 1978. Systematics and evolution of Garrya. Contributions to the Gray Herbarium of Harvard University 209:1—104. EDGAR, R. C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32:1792—1797. FELSENSTEIN, J. 1985. Phylogenies and the comparative method. American Naturalist 125:1—15. HALLOCK, F. A. 1930. The relationship of Garrya. The development of the flowers and seeds of Garrya and its bearing on the phylogenetic position of the genus. Annals of Botany 176:771—812. LISTON, A. 2003. A new interpretation of floral morphology in Garrya (Garryaceae). Taxon 52:271-276. MADDISON, W. P. AND L. L. KNOWLES. 2006. Inferring phylogeny despite incomplete lineage sorting. Systematic Biology 55:21—30. Munz, P. A. AND D. D. KEck. 1968. A California flora and supplement. University of California Press, Berkeley, CA. NYLANDER, J. A. A. 2004. MrModeltest, v2. Program distributed by the author, Evolutionary Biology Centre, Uppsala University, Uppsala, Sweden. RONQUIST, F. AND J. P. HUELSENBECK. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19:1572—1574. SOLTIS, D. E., P. S. SoLTIs, M. W. CHASE, M. E. Mort, D. C. ALBACH, M. ZANIS, V. SAVOLAINEN, W. H. HAHN, S. B. Hoot, M. F. FAy, M. AXTELL, S. M. SWENSEN, L. M. PRICE, W. J. KRESS, K. C. NIXON, AND J. S. FARRIS. 2000. Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Botanical Journal of the Linnean Society 133:381-461. SWOFFORD, D. L. 2000. PAUP*. Phylogenetic analysis using parsimony (*and other methods). Version 4.01b10, Sinauer Associates, Sunderland, MA. TABERLET, P., G. LUDOvIC, G. PAUTOU, AND J. BOUVET. 1991. Universal primers for amplification of three non-coding regions of chloroplast DNA. Plant Molecular Biology 17:1105—1109. WHITE, T. J., T. BRUNS, S. LEE, AND J. TAYLOR. 1990. Amplifications and direct sequencing of fungal ribosomal RNA genes for phylogenetics. Pp. 315— 322 in M. A. Innis, D. Gelfand, J. J. Sninsky, and T. J. White (eds.), PCR protocols: a guide to methods and applications. San Diego: Academic Press, San Diego, CA. ZWICKL, D. J. 2006. Genetic algorithm approaches for the phylogenetic analysis of large biological se- quence datasets under the maximum likelihood criterion. Ph.D. dissertation, The University of Texas, Austin, TX. 2011] APPENDIX | SAMPLED INDIVIDUALS OF GARRYA AND AUCUBA Collector and number followed by description of locality. All specimens deposited in the Duke University Herbarium (DUKE). Aucuba japonica Thunb.—D.O. Burge 363, City of Chico, 1469 Humboldt Rd, Butte Co., CA. Garrya buxifolia Gray—D.O. Burge 1160, Swede Creek water- shed, roadside on FR 2524 (road to Spalding Mill), Josephine Co., OR. G. elliptica Douglas ex Lindl.— D.O. Burge 382, Seaside, eastern terminus of Kimball Avenue near Fort Ord Military Reservation, Monterey Co., CA; D.O. Burge 386, Mount Tamalpais, roadside on East Ridgecrest Boulevard, near Middle Peak, Marin Co., CA. G. flavescens S. Watson —D.O. Burge 370, Ball Mountain, western slope, along Caliente Bodfish Rd, Kern Co., CA; D.O. Burge 419, Wilson Mountain, Wilson Mountain Trail, Yavapai Co., AZ; D.O. Burge 1036, Cerro Bola, eastern slope, Baja California, Mexico. G. fremontii Torr.—D.O. Burge 353, Doe Mill Ridge (ridge between Butte Creek and Little Chico BURGE: PHYLOGENETICS OF GARRYA aes) Creek), Butte Co., CA; D.O. Burge 362, Trinity River canyon, Poison Gulch, Humboldt Co., CA; D.O. Burge 1148, North Fork Tuolumne River watershed, Bald Mountain, Tuolumne Co., CA. G. glaberrima Wan- gerin—D.O. Burge 1025, Cerro Juarez, near summit, Hidalgo, Mexico; D.O. Burge 1216, Cerro El Potosi, eastern slope, Nuevo Leon, Mexico; D.O. Burge 1225, Sierra El Pedregoso, Tamaulipas, Mexico. G. grisea Wiggins—D.O. Burge 778, Sierra San Pedro Martir, Baja California, Mexico. G. laurifolia Benth.—D.O. Burge 1218, Cerro El Potosi, eastern slope, Nuevo Leon, Mexico; D.O. Burge 1252, Cascada de Basaseachi area, Chihuahua, Mexico. G. lindheimeri Torr.—D.O. Burge 750, City of Austin, Mayfield Park and Nature Preserve, Travis Co., TX. G. ovata Benth.—D.O. Burge 1221, Sierra Los Soldados, Nuevo Leon, Mexico. G. veatchii Kellogg—D.O. Burge 378, Cuesta Ridge, San Luis Obispo Co., CA; D.O. Burge 1041, Isla Cedros, N slope of Cerro Redondo, Baja California, Mexico. G. wrightii Torr.—D.O. Burge 934, Santa Catalina Mountains, Pima Co., AZ; D.O. Burge 1239, Sierra de Coneto, western slope, Durango, Mexico; D.O. Burge 1253, Cascada de Basaseachi area, Chihuahua, Mexico. MADRONO, Vol. 58, No. 4, pp. 256-257, 2011 LECTOTYPIFICATION OF ARCTOSTAPHYLOS HOOVERI (ERICACEAE) DAVID J. KEIL Biological Sciences Department, California Polytechnic State University, San Luis Obispo, CA 93407 dkeil@calpoly.edu ABSTRACT The holotype of Arctostaphylos hooveri P. V. Wells, cited as having been deposited in the Robert F. Hoover Herbarium, California Polytechnic State University (OBI), apparently is not extant. The isotype deposited in the University Herbarium (UC) at the University of California, Berkeley (/Toover & Wells 1960 — UC 1218855) is designated as the lectotype, and the isotype in the California Academy of Science Herbarium (CAS 423565) is thereby an isolectotype. Hoover 8520 (OBI 175682) is recognized as a paratype. Key Words: Arctostaphylos hooveri, Ericaceae, lectotypification. Wells (1961) described a new species, Arcto- staphylos hooveri, from Monterey Co., CA. In his introduction Wells stated: “In April, 1960, Dr. Robert F. Hoover collected flowering specimens of a Manzanita near the summit of Nacimiento Pass, Monterey Co., California, which appeared to be a variant close to Arctostaphylos andersonii A. Gray. The writer visited the locality on May 1, and made additional collections and observa- tions.”” Apparently Hoover and Wells traveled together on this second trip. In the type citation Wells stated that “‘the [holo]type, Hoover & Wells 1960, collected on May 1, 1960, is in the herbarium at California Polytechnic College, San Luis Obispo [OBI]. Isotypes are at the California Academy of Sciences [CAS] and University of California, Berkeley [UC]. The label on the UC isotype was typed onto a University of California label form and_ bears the following data: California. Monterey Co.: “Tall erect shrub to over 12 feet high, with well- defined trunk lacking a basal burl. Roadside clearings in forest of broad sclerophylls and local Pinus ponderosa; on shallow soils over gneissic bedrock; along road leading south from Naci- miento Pass, altitude ca. 3000 feet.” The number /960 is not Hoover’s collection number; Hoover 1960 was a 1937 Lupinus colle- ction from Tuolumne Co., CA. Nor was it Wells’s number; according to collection records in the Consortium of California Herbaria data- base (CCH), Well’s collection numbers between April 12 and June 30 of 1960 ranged from 43 to 75. The number /960 most likely represents the year of the collection. A specimen was located among the OBI types labeled Arctostaphylos hooveri Wells sp. nov. (OBI 75682). The words ““TYPE SHEET?!” are printed in pencil in an unknown hand directly on the mounting paper next to the label. The collecting data were typed onto one of Hoover’s personal collecting labels (which actually bears a pre-printed San Luis Obispo Co. heading): ““Monterey County. In roadside chaparral near summit of Nacimiento Pass. Erect, without burl. April 1960.” The collection number 8520 was hand-written in ink, probably by Hoover, as is staphylos hooveri Wells’. The herbarium identi- fication “HERBARIUM of California State Polytechnic College, San Luis Obispo, Califor- nia’’ was stamped onto the mounting paper at an unknown date, presumably before 1972 when Cal Poly became a university. Hoover’s collection notebook records for 1960 (deposited in OBI) do not record either the April 1960 excursion or a collecting trip with P. V. Wells in May. Hoover did not record trips in which he made no numbered collections. On April 9, 1960 Hoover collected numbers 85/7—8520 along the coast of San Luis Obispo Co. from the vicinity of Lion Rock to the mouth of Coon Creek (localities now part of the Pacific Gas and Electric Company’s Field Ranch holdings north of Diablo Canyon Nuclear Power Plant). According to Hoover’s notes, the collec- tion number 8520 is a Ceanothus from the mouth of Coon Creek. The notebook entry (in ink) is followed by a penciled *“‘Molded.”” Examination of the Hoover 8520 Ceanothus specimen in OBI (Ceanothus thyrsiflorus var. griseus) reveals that it did, indeed, suffer mold damage during drying, but it was nevertheless processed. The next entry in Hoover’s notebook, 852/, was an Allium specimen collected on May 13, 1960 from Rinconada Mine in San Luis Obispo County. It would appear that Hoover retroactively assigned the collection number 8520 to the Mon- terey Co. Arctostaphylos specimen, thinking that he would discard the mold-damaged Ceanothus 2011] specimen to which he had originally assigned that number. But both were mounted and are now deposited in OBI. No specimen of Hoover and Wells 1960 has been located in OBI. All California Arctostaph- ylos specimens in OBI have been databased, and the records have been submitted to CCH. As far as I am able to determine the holotype of A. hooveri is not extant. Accordingly, I am _ here designating the isotype deposited in UC (Hoover & Wells 1960 — UC 1218855) as the lectotype; the isotype in CAS (CAS 423565) 1s thereby an KEIL: LECTOTYPIFICATION OF ARCTOSTAPHYLOS HOOVERI 25) isolectotype. Hoover 8520 (OBI 75682) is appar- ently the April 1960 specimen mentioned by Wells (1961) and is here recognized as a paratype of A. hooveri; no isoparatypes are cited in CCH. LITERATURE CITED CONSORTIUM OF CALIFORNIA HERBARIA. Website http://ucjeps.berkeley.edu/consortium/ [accessed 03 Feb 2012]. WELLS, P. V. 1961. A new manzanita from the Santa Lucia Range, California. Leaflets of Western Botany 9:152—153. MADRONO, Vol. 58, No. 4, pp. 258-266, 2011 CALYPTRIDIUM PARRYI VAR. MARTIRENSE (MONTIACEAE), A NEW TAXON ENDEMIC TO THE SIERRA DE SAN PEDRO MARTIR, BAJA CALIFORNIA, MEXICO C. MATT GUILLIAMS! AND MICHAEL G. SIMPSON Department of Biology, San Diego State University, San Diego, CA 92182, USA matt_g@berkeley.edu JON P. REBMAN San Diego Natural History Museum, P.O. Box 121390, San Diego, CA 92112-1390 ABSTRACT Calyptridium parryi var. martirense is described as new. Here we present quantitative measurements and statistical analyses of a number of morphological features that demonstrate the distinctiveness of this new taxon. The new variety differs from the others in having shorter fruits (3.1-4.1 mm) and a correspondingly smaller fruit length to sepal length ratio (1.0—1.4). The capsule is also the widest (1.4— 2.2 mm) and the sepals the longest (2.4-3.9 mm) of any other C. parryi variety. Calyptridium parryi var. martirense is currently known only from high elevation locations (1900—2630 m) in the Sierra de San Pedro Martir, Baja California, Mexico. Key Words: Baja California, Calyptridium, endemism, Montiaceae, Sierra de San Pedro Martir. Calyptridium (Montiaceae, sensu APG III 2009) is a genus of annual and perennial herbs comprising nine species and 11 total taxa, including varieties. (See Simpson et al. 2010 for a review of the taxonomy of the genus and associated literature.) The genus has been the subject of a recent master’s thesis (Guilliams 2009), and a morphological analysis of the currently recognized varieties of C. parryi A. Gray plus the morphologically similar C. mon- andrum Nutt. (Simpson et al. 2010). In that latter study we detected significant morphological differences between C. parryi specimens from the Sierra de San Pedro Martir of Baja Califor- nia, Mexico, and all other specimens of C. parryi included in the analysis, although only five samples of the former were measured. These specimens from the Sierra de San Pedro Martir had previously been identified as either C. monandrum or C. parryi var. nevadense J. T. Howell. Here we provide quantitative data from all known specimens of this annual Calyptridium from the Sierra de San Pedro Martir, confirming the morphological pattern previously reported by Simpson et al. (2010). We conclude that these collections from the Sierra de San Pedro Martir represent a new taxon, which we recognize as a variety of C. parryi. MATERIALS AND METHODS This study includes quantitative analyses of a total of 85 specimens of C. parryi varieties (var. ‘Present address: Department of Integrative Biology, University of California Berkeley, 1001 Valley Life Sciences Building #2465, Berkeley, CA 94720-2465. hesseae J. H. Thomas, var. nevadense, var. parryi, and the variety described here) from 10 herbaria (ARIZ, ASU, CAS-DS, JEPS, RSA, SD, SDSU, UC, UCR, UTC). In addition, we included 12 specimens of C. arizonicum (J. T. Howell) M. G. Simpson, M. Silveira & Guilliams and 13 of C. monandrum, taxa that we formerly recognized as distinct from C. parryi (Simpson et al. 2010), so that our previous taxonomic hypothesis could be evaluated with this expanded dataset. The data matrix from the previous study was broadened to include complete measurements of all remaining C. parryi specimens from Baja California, Mex- ico, a total of 11 additional specimens. Only one specimen was excluded from the analysis (Vasek s.n., UCR 15269), as the individuals on this sheet do not bear mature fruits. Vouchers of all specimens of Calyptridium arizonicum, C. mon- andrum, and C. parryi vars. hesseae, nevadense, and parryi used in the current analysis are the same as those cited in Appendix | of Simpson et al. (2010); vouchers of the Sierra de San Pedro Martir populations of C. parryi used in the current analysis are equivalent to the list of types below, minus the Vasek s.n. specimen. Characters measured and methods of data acquisition were identical to Simpson et al. (2010). In brief, several seed measurements were taken, including the seed sagittal diameter (including notch length), seed transverse diame- ter, seed notch length, and distance from the perimeter to the central-most extent of papillae along both the transverse and sagittal planes (these last two transformed as percentage dis- tance from perimeter to center of the seed). In addition to seed characters, we measured fruit 2011] GUILLIAMS ET AL.: A NEW VARIETY OF CALYPTRIDIUM PARRYI 259 TABLE 1. DESCRIPTIVE STATISTICS FOR CALYPTRIDIUM TAXA EXAMINED IN THIS STUDY. Taxon acronyms: C. arizonicum = ARI; C. monandrum = MON; C. parryi var. hesseae = HES; C. parryi var. martirense = MAR; C. parryi var. nevadense = NEV; C. parryi var. parryi = PAR. Values provided are the mean, minimum/maximum values, and the standard deviation. Asterisks (* = P < 0.05; ** = P < 0.01) and plus signs (+ = P < 0.05; ++ = P < 0.01) indicate when a taxon is statistically different from all other taxa or from other members of Calyptridium parryi, respectively, in Anova/Tukey post-hoc tests of significance. Seed sagittal Taxon n diameter (mm) ARI 12 0.759** 0.705/0.803 0.030 MON 13 0.615 0.538/0.692 0.055 HES 16 0.542** 0.483/0.576 0.025 MAR 16 0.635 0.568/0.723 0.051 NEV 33 0.675 0.578/7.58 0.041 PAR 20 0.663 0.547/0.740 0.052 length, maximum fruit width, sepal blade length, sepal width at widest point, scarious sepal margin width (measured on one side only, the one most intact), and distance from the sepal blade base to the point of greatest sepal width. One ratio character was created and examined: ratio of fruit length to sepal length. See Simpson et al. (2010) for a diagrammatic representation of these measurements. Most statistical analyses used in Simpson et al. (2010) were repeated with the present data set, although here we omit the principal components analysis for brevity. The morphological charac- ters that appeared important in distinguishing between the Sierra de San Pedro Martir speci- mens of C. parryi and the three recognized varieties were evaluated for statistically signifi- cant differences by taxon using analysis of variance (ANOVA), with multiple comparisons made between the taxa for each character using the Tukey post hoc test. All statistical analyses were performed in SYSTAT, Version 11 (Systat Software, Inc., San Jose CA; http://www.systat. com). RESULTS Basic descriptive statistics for the variables measured are given in Table 1. Importantly, these data continue to support Calyptridium arizonicum and C. monandrum as_ separate taxonomic entities, with strong differences in fruit length, seed size, and seed sculpturing in the former, and sepal length, fruit width, and fruit length to sepal length ratio in the latter. With respect to C. parryi, seed size, seed papillation, fruit length, fruit width, width of the sepal scarious margin, and the fruit length to sepal length ratio appear to be among the more important of the morphological traits Seed notch length (mm) Seed transverse diameter (mm) 0.750* 0.705/0.782 0.028 0.022 0.016/0.031 0.005 0.634 0.553/0.717 0.056 0.018 0.010/0.024 0.004 0.538** 0.495/0.587 0.025 0.023 0.016/0.031 0.004 0.669 0.598/0.756 0.051 0.037 0.019/0.056 0.009 0.703 0.640/810 0.036 0.045 0.010/0.354 0.056 0.691 0.570/0.765 0.050 0.032 0.022/0.046 0.007 distinguishing the varieties from one another. These differences between varieties /hesseae, nevadense, and parryi are discussed at length in Simpson et al. (2010) and will not be repeated here. The remainder of this section will focus on those quantitative characters important for evaluating the distinctiveness of C. parryi ‘“‘mar- tirense.”’ Fig. 1A—C shows boxplots for some of these variables, with asterisks denoting when a given taxon was found to be statistically different from all other taxa and plus signs denoting when a subgroup (variety or the “‘martirense’’ speci- mens) of C. parryi was statistically different from all other subgroups of C. parryi, using the Tukey HSD post hoc test (described below). For C. parryi “martirense’’, the fruit length (3.1—4.1 mm, mean = 3.7 mm), fruit width (1.42.2 mm, mean 1.9 mm), fruit length to sepal length ratio (1.0— 1.4, mean = 1.2), sepal length (2.4-3.9 mm, mean 3.2 mm), and sepal width (2.4-3.8 mm, mean 3.2 mm) appear to be the most distinctive quantitative traits with respect to other C. parryi varieties (Table 1, Fig. 1). The ANOVAs and Tukey HSD post hoc tests of these variables confirm that the varieties of C. parryi, including C. parryi “‘martirense,” are significantly different from one another (Table 1). Among all studied taxa, Calyptridium parryi “‘martirense” has a significantly smaller fruit length (Fig. 1A), and a significantly smaller fruit length to sepal length ratio (Fig. 1B). Among the C. parryi varieties, ““martirense”’ has a significantly larger fruit width (Fig. 1C), a significantly larger sepal length (Fig. 1D), and an intermediate but significantly different sepal width (Table 1). DISCUSSION This study expands upon the morphometric analysis of Simpson et al. (2010) by including all 260 MADRONO [Vol. 58 TABLE 1. EXTENDED. Seed papillation Seed papillation Fruit Fruit Taxon % (sagittal) % (transverse) length (mm) width (mm) ARI 0** 0/0 0 00** 0/0 0 6.689** 5.817/ 1.742 1.346/ 7.775 0.650 1.943 0.169 MON 34.910 28.144/ 35.922 29.184/ 4.384 3.780/ 0.894** 0.790/ 40.746 4.307 92.527.5.812 5.044 0.442 1.100 0.076 HES 31.543 26.821/ 33.988 27.662/ 4.468 4.110/ 1.449 1.110/ 35.032 2.235 38.912 2.954 4.691 0.200 2.018 0.273 MAR 39.529 30.082/ 38.024 31.117/ 3.676** 3.067/ 1.898++ 1.435/ 51.657 6.077 46.712 4.469 4.117 0.372 2.243 0.231 NEV 35.290 25.316/ 34.208 25.574/ 4.689 3.830/ 1.504 1.080/ 54.351 6.845 52.043 6.244 5.745 0.533 1.927 0.172 PAR 99.720** 94.396/ 99.668** 93.362/ 4.567 2.913/ 1.577 1.340/ 100 1.253 100 1.484 5.260 0.595 1.863 0.150 known accessions of C. parryi “‘martirense’”’ from the Sierra de San Pedro Martir. The Simpson et al. (2010) study focused on examining differ- ences between C. arizonicum and C. parryi, and thus only five accessions of C. parryi “‘martirense”’ were measured and included. Nevertheless, we found patterns in the morphological data that suggested that C. parryi “‘martirense”’ was mor- phologically distinct from the other varieties of C. parryi. The present study with expanded sampling confirms our previous results, showing that C. parryi “‘martirense” differs from other C. parryi varieties in a number of morphological features, the most notable of which are fruit length and fruit length to sepal length ratio. While the differences between the C. parryi varieties are minute, in some cases requiring a dissecting microscope to adequately quantify (e.g., seed size, seed papillation), they are remarkably consistent and statistically significant. The demonstrated morphological distinctive- ness of these taxa, coupled with largely non- overlapping distributions, provide compelling justification for retaining the current varieties of C. parryi, as well as for recognizing C. parryi ‘““martirense” as a new taxon, using a taxonomic species concept (Cronquist 1978, 1988). Under this species concept, also known as the morpho- logical species concept, species are circumscribed based on the discontinuity of morphological features with respect to other morphologically similar species. Varieties and subspecies are typically considered to be taxonomic entities that show morphological differences from one anoth- er, but the feature or features that make them different have some intergradation. This present study also supports previous taxonomic hypotheses regarding the distinctive- ness of C. arizonicum and C. monandrum, taxa that are morphologically similar to the C. parryi varieties. Calyptridium parryi var. martirense Guilliams, M. G. Simpson, & Rebman, var. nov. (Figs. 2, 3).—Type: MEXICO, BAJA CALIFORNIA, Sierra de San Pedro Martir, S of Vallecitos in wet, open meadow near Cerro la Botella Azul, Pinus jeffreyi, Abies concolor, Linanthus melingii, granitic sand, 2590 meters elev., 30.97222°N, 115.43722°W, 28 June 1998, J. P. Rebman 5407 (holotype: SD; isotypes: BCMEX, MEXU, RSA, UC, UCR). Calyptridium parryi var. martirense differt a C. parryi vars. hesseae, nevadense, et parryi fructibus brevioribus (3.1—4.1 mm) et latioris (1.4—2.2 mm), sepalis longioris (2.4-3.9 mm), et proportione parve longitudinis fructus ad sepali (1.0—1.4). Calyptridium parryi var. martirense differs from C. parryi vars. hesseae, nevadense, and parryi by a shorter (3.14.1 mm) and wider (1.4-2.2 mm) fruit, by a longer sepal (2.4— 3.9 mm), and by a small fruit to sepal length ratio (1.0—1.4). Plant an annual or possibly biennial herb. Root a single taproot. Stems prostrate, with several (5— 13) primary branches radiating from center, primary branches up to 13 cm long, each terminating in an inflorescence unit and either unbranched or (more typically) bearing several lateral secondary and tertiary branches, each of these with a terminal inflorescence unit; stems becoming pink at maturity in some plants. Leaves simple, spiral, exstipulate, basal and cauline in position, sessile from narrow, attenuate base (appearing petiolate), often proximally condupli- cate-canaliculate and forming membranous, sheath-like margins at point of attachment, margin entire, apex acute-rounded, mucronulate, glabrous, somewhat succulent, only midrib vein apparent; basal leaves forming compact, outer rosette when immature, often caducous when mature, mature leaves spatulate, up to 5 cm long, 5 mm wide at widest region near apex; cauline 2011] GUILLIAMS ET AL.: A NEW VARIETY OF CALYPTRIDIUM PARRYI 261 TABLE |. EXTENDED. Sepal blade Sepal Sepal scarious Sepal base to Fruit length: Taxon length (mm) width (mm) margin (mm) widest (mm) sepal length ARI 3.090 2.756/ 3.942 3.416/ 0.415 0.345/ 0.885 0.709/ 2.173 1.885/ 3.783 0.311 4.725 0.395 0.470 0.042 1.066 0.118 2.455 0.169 MON 1.591** 1.349/ 1.419 1.238/ 0.161 0.051/ 0.633* 0.459/ 2.781** 2.398/ 2.027 0.203 17310152 0.248 0.059 1.023 0.159 3.418 0.342 HES 2.297* 1.674/ 1.838+4+- 1.353/ 0.1604++ 0.053/ 0.894 0.418/ 1.967+ 1.667/ 2.553 0.269 2.524 0.299 0.237 0.051 1.229 0.196 2.563 0.217 MAR 3.186+4+ 2.437/ 3.165+4+4 2.412/ 0.447 0.348/ 0.981 0.719/ T.173** 1.025/ 3.918 0.412 3.796 0.414 0.687 0.089 1.239 0.164 1.446 0.102 NEV 272 1107 3.715+ 2.248/ 0.634** 0.339/ 1.074 0.590/ 1.734 1.157/ 3.827 0.500 5.253 0.800 1.048 0.169 1:3220.257 2.595 0.312 PAR 2.702 2.150/ 2.765 2.076/ 0.414 0.265/ 1.103 0.766/ 1.705 1.117/ 3.272 0.346 3.570 0.375 0.629 0.094 1.448 0.199 2.172 0.239 leaves spatulate to oblanceolate, decreasing in size toward apex, becoming pink at maturity in some plants. Inflorescence units terminal on primary or lateral branches, bracteate, bracts subtending axes usually photosynthetic, short oblanceolate, mu- cronulate to mucronate; bractlets present at base of and along axis of inflorescence unit scarious, triangular to ovate, acuminate; inflorescence unit a secund, monochasial cyme, the flower-contain- ing part of units up to 3 cm long at maturity. Flowers perfect, bracteate, subsessile. Flower bracts positionally displaced from flowers, ca. 2 mm long, white-scarious, sessile, lance-ovate to ovate-deltate, rounded to cordate, entire, caudate, glabrous. Perianth biseriate. Calyx aposepalous, green with whitish, scarious margin, glabrous. Sepals two, unequal, at flowering stage adaxial sepal widely ovate, ca. 3 mm long, 2.5 mm wide, strongly involute distally, overlapped by abaxial sepal; abaxial sepal widely orbicular, in flower ca. 3 mm long, 3.5 mm wide, with scarious margins ca. 0.5 mm wide, mostly widely ovate to orbicular; sepals accrescent at fruiting stage, abaxial sepal at fruiting stage widely orbicular, 2.4-3.9 mm long, 2.4-3.8 mm wide, cuneate at base, reniform just above base, entire or irregularly lobed, undulate when dried, apically rounded, mostly ternately veined from base, veins anastomosing, prominent when dried, margins scarious, scarious region 0.3— 0.7 wide on each side; abaxial sepal appressed to fruit. Corolla apopetalous, actinomorphic. Petals whitish, four (rarely five), quincuncial, oblong, apically rounded, slightly cup-shaped (concave toward central axis), ca. 2 mm long, 1 mm wide. Stamens three, uniseriate, apostemonous, fila- mentous, whorled, inserted. Filaments terete, ca. 0.5 mm long. Anthers basifixed, dithecal, longitu- dinal, introrse, ca. 0.5 mm long, elliptic, thecae parallel. Pollen yellow. Gynoecium syncarpous, hypogynous. Ovary superior, ca. 1 mm long, globose, 3-lobed, glabrous. Style one, terminal, terete, ca. 0.3 mm long. Stigmas 3, narrowly oblong, slightly twisted, ca. 0.2 mm long, papil- late. Nectaries not observed. Carpels 3. Locule 1. Placentation free-central. Ovules approximately 14 per ovary. Fruit a 2-valved, tan, oblong capsule, flattened perpendicular to inflorescence axis, 3.0-4.1 mm long and 1.4-2.2 mm wide at maturity apically calyptrate from persistent, detached corolla. Seeds ca. 6—10 per fruit, black, discoid, with marginal notch, 0.6—0.7 mm in diameter, glabrous, smooth in center, papillate along margin (width of papillate region ca. 0.1— 0.2 mm), arranged in 2 rows in fruit. Calyptridium parryi var. martirense 1s currently known only from high elevation (1900-2630 m) locations in the Sierra de San Pedro Martir of Baja California, Mexico. It is found in usually sandy soil and/or soil and rocks of granitic origin, sometimes fine clay soil, and typically in open habitats near creeks/streams or in wet or dry, open meadows or forest understory of mixed forests of Pinus jeffreyi, Abies concolor, Populus tremuloides, and occasionally Hesperocyparis montana (=Cupressus m.; Callitropsis m.) with mixed shrub and herb associates. The Sierra de San Pedro Martir is a floristically diverse region of great botanical importance, having a natural fire regime and being the southern-most limit of several montane plant species of the California Floristic Province (Riemann and Ezcurra 2007; Thorne et al. 2010). The higher elevations comprise the Parque Nacional Sierra de San Pedro Martir, established in 1947. Thorne et al. (2010) reviewed the vascular plant flora of the “high” Sierra de San Pedro Martir, defined as being greater than 1800 meters in elevation. These authors cited 453 species native to this region. Of these taxa, 23 species and one variety are endemic to the Sierra de San Pedro Martir, slightly over 5%. To this we add another variety, increasing the endemic flora of this interesting region. Note that in addition to Calyptridium parryi var. martirense, Thorne et al. N ON i) 6 E E = =) = 9 CF) wl = = te ica 2.0 Fruit Width (mm) pe 1.0 0.5 Fic. 1. MADRONO [Vol. 58 Fruit Length : Sepal Length Sepal Length (mm) Box plots of single characters. A. Fruit length (mm). B. Fruit length to sepal length ratio. C. Fruit width (mm). D. Sepal length (mm). Note: box plots show median (middle horizontal line), first and third quartiles (lower and upper horizontal lines, respectively), and the range of the data outside the first and third quartiles (vertical lines); x = outliers; C. parryi ““‘martirense” = San Pedro Martir Mountains populations of Calyptridium parryi. Statistical difference between a given taxon and all other taxa (via ANOVA Tukey post hoc test) is indicated as: ** = P< 0.01; * = P < 0.05; that between a given member of C. parryi and all other C. parryi taxa only is indicated as: ++ = P < 0.01; + = P < 0.05. (2010) cite two other species of the genus as occurring in the Sierra de San Pedro Martir, C. monandrum Nutt. and C. monospermum Greene. Based upon the morphological features presented in this paper, C. parryi var. martirense is easily distinguished from C. monandrum, and neither of these annual taxa are likely to be confused with the more robust perennial species, C. monospermum. Calyptridium parryi var. martirense flowers from June to as late as early August and develops mature fruits from June to as late as October. The derivation of the varietal epithet, martir- ense, 18 after the Sierra de San Pedro Martir, (“mountain of Saint Peter the martyr’’), to which this variety is endemic. We suggest the Sierra de San Pedro Martir Calyptridium as the common name for the taxon. Paratypes (see Fig. 4 for map of localities): MEXICO, BAJA CALIFORNIA. Sierra San Pedro Martir, along sandy roadside in forest of Jeffrey pine and white fir, 31.03676°N, 115.473°W (lat/long. estimated from locality data), 2590 m elev., 24 July 1975, Almeda 2582 (CAS 611882); Sierra San Pedro Martir, mead- ows along road heading S of Vallecitos towards La Encantada, base of Cerro Botella Azul, surrounding forest of Pinus jeffreyi, Abies con- color, Populus tremuloides, prostrate, succulent | Phas : _ > 8 eae | “ae — Pea 28. Bes capsule i valve scarious margin FIG. 2. A-C. Photographs of Calyptridium parryi var. GUILLIAMS ET AL.: A NEW VARIETY OF CALYPTRIDIUM PARRYI 263 fruite™ cep: 7. A ipsule) ot ! v ‘ef “ ae | ae 4:44 martirense in native habitat. A—B. Plant habit (Rebman 15994), at early stage (A) and mature stage (B). C. Close-up of fruit with abaxial sepal (Rebman 16055). D. Close-up of fruit and abaxial sepal from herbarium specimen (Moran 24462). E. Seed in face (left) and side (right) views, showing notch, papillate margin, and smooth/shiny central region (Rebman 5407, type specimen). annual, common in sandy forest understory and drier, sandy areas of meadow, 30.9907°N, 115.4403°W, 2510 m elev., (lat/long. and eleva- tion estimated from locality data), 19 July 1988, Boyd 2645 (RSA _ 519524); Sierra San Pedro Martir, canyon at base of Cerro Botella Azul, at end of road leading S of Vallecitos, surround- ing forest of Pinus jeffreyi, Abies concolor, Populus tremuloides, Cupressus montana, locally common, prostrate, succulent annual, infrequent in sandy openings, 30.99071°N, 115.44030°W, 2509 m elev., (lat./long. and elevation estimated from locality data), 19 July 1988, Boyd 2706 (RSA 519494); Yerba Buena, 31.000°N, 115.450°W, 2475 m elev., abundant in sandy soil, 16 August 1967, Moran 14185 (RSA 225157); La Vibora, Arroyo la Grulla 4 km SW of La Grulla., occasional in dry sand by stream, 30.867°N, 115.508°W, 1900 m elev., 10 August 1977, Moran 24462 (SD 97873); Rancho Viejo, 30.900°N, 115.483°W, 2050 m elev., fairly common in dry, sandy meadow, 11 August 1977, Moran 24489 (SD 97766); a bit W of Vallecitos along trail from Prado del Corona to La Providencia, N end of high Sierra San Pedro Martir, 30.98570°N, 115.51824°W, (lat./long. estimated from locality data), 2350 m elev., sandy, sunny flat, small, wide streambed; aspens, Salix, Urtica nearby, rosettes to 7 in. across, frequent, early to mid-bloom, 07 June 1962, Olmsted 4603 (RSA 170797); Sierra San Pedro Martir, La Encantada meadow, S of Los Llanitos, leaves succulent, fruit shorter than C. monandrum, sepals round with small margins, 30.91667°N, 115.40000°W, 2200 m elev., 27 June 264 MADRONO [Vol. 58 S.FAWCET 2010 Fic. 3. Line drawings (by Susan Fawcett) of type specimen of Calyptridium parryi var. martirense, from type specimen (Rebman 5407). A. Whole plant. B. Inflorescence unit. C. Fruit with adaxial sepal. D. Calyptra, removed from fruit. 2011] GUILLIAMS ET AL.: A NEW VARIETY OF CALYPTRIDIUM PARRYI Sierra de San Pedro Martir Observatory eS Y ~ 1,600 m Cerro Picacho del Diablo cs ® San Pedro Martir Fic. 4. Map showing the geographic distribution of the 17 known collections (circles) of Calyptridium parryi var. martirense from Sierra de San Pedro Martir. Locality indicated with ““X”’ is that of the type specimen, Rebman 5407. 1996, Rebman 3295 (SD 142900); Sierra de San Pedro Martir, Vallecitos area, just S of the main road to the Observatory, Pinus jeffreyi, Eriogo- num wrightii var. oresbium, granitic substrates, annual, note fruit differences from C. monandrum, 31.02333°N, 115.47000°W, 2415 m elev., 30 July 1997, J. Rebman 4174 (SD 142899); Sierra San Pedro Martir, La Tasajera region, SW of Observatory, approx. 7 mi S of the Observatory Rd, Pinus jeffreyi, Abies concolor, Populus tremu- loides, granite rocks and soil, 30.94389°N, 115.49722°W, 2285 m elev., 15 September 1998, J. Rebman 5579 (SD 145561); Sierra San Pedro Martir, SE of Vallecitos and approx. 3 mi S of the Observatory, along the highest ridge en route to Pedro’s Dome, conifer forest with Pinus jeffreyi, 266 Abies concolor, Eriogonum wrightii var. oresbium, Philadelphus microphyllus, and Callitropsis mon- tana, mostly granitic substrates, prostrate, annual, rare, 31.008°N, 115.436°W, 2630 m elev., 30 September 2008, Rebman 15994 (SD 191485); Sierra San Pedro Martir, meadow along the road to Tasajera approx. 0.75 mi S of the Observatory Rd in Vallecitos, meadow surrounded by mixed conifer forest with Populus tremuloides, Pinus contorta, Pinus jeffreyi, Achillea millefolium, Calyptridium parryi, and Xanthisma_ wigginsii, granitic substrates, 31.0042°N, 115.4925°W, 2465 m elev., 01 October 2008, Rebman 16055 (SD 191486); Sierra San Pedro Martir, “Corral Meadow” 7.5 km NW (340 degrees) of the observatory, decomposed granite soil on slopes, fine clay in meadow, slopes with mixed conifer forest of Abies concolor and Pinus jeffreyi, meadow with Juncus, Poa, Carex, etc., a fairly common annual on open flats, succulent leaves, 31.11250°N, 115.49722°W, 2520 m elev., 16 June MADRONO [Vol. 58 1988, Sanders 7921 (UCR 52532); Vallecitos, near road to Observatory and camp-ground, open, sandy meadow and stream, dry, sandy soil of meadow, succulent, spreading annual, 31.033°N, 115.467°W, 2430 m elev., 18 June 1985, Thorne 60834 (RSA 346089); Sierra San Pedro Martir, study area 3, on observatory road 7.5 mi above Parque Nacional San Pedro Martir boundary (=entrance station?), Jeffrey pine forest with Blepharoneuron tricholepis, Aristida, Hymenopap- pus filifolius, Machaeranthera wigginsii, Muhlen- bergia minutissima, Draba corrugata, Ipomopsis effusa, Potentilla wheeleri, Monardella macrantha, Gayophytum diffusum, Ophiocephalus angustifo- lius, Eriogonum hastatum, etc., 31.033°N, 115.467°W, 2410 m elev., 17 October 1976, Vasek, s.n. (UCR 15269); Vallecitos, Sierra San Pedro Martir, 31.000°N, 115.467°W, 2475 m elev., sandy ground near creek, 09 August 1969, Witham 384 (SD 74689). A Revised Key of the Varieties of Calyptridium parryi and Closely Related Species Fruit at maturity gen. <] mm wide, >2.5 = longer than abaxial fruiting sepal; abaxial fruiting sepal <2 mim long: widespread... ..« «44. 4.64644 +0644 C. monandrum Fruit at maturity gen. >1 mm wide, <2.5 X longer than abaxial fruiting sepal; abaxial fruiting sepal 1.7— 4 mm long; restricted in range Seeds completely smooth, lacking papillae; fruit gen. 5.8-8 mm long................ C. arizonicum Seeds not completely smooth, with papillae at least along the margin; fruit 3—5.7 mm long. Seeds with papillae throughout.......... Seeds with papillae only on the margin C. parryi var. parryi Fruit ca. 3-4 mm long, =1.5 X longer than abaxial fruiting sepal; Sierra de San Pedro Martir, Baja California, Mexico............ ee eee ee eee Orr ie C. parryi var. martirense Fruit 3.8—5.7 mm long, 1.5(1.1)—2.5 X longer than abaxial fruiting sepal; mostly California and Nevada (rarely Utah, Arizona), USA Seeds >0.6 mm in diameter; abaxial fruiting sepal reniform with a wide scarious margin; mostly Sierra Nevada and Panamint ranges, eastern California to western Nevada (rarely Ulan AniZona pa. 7 2 ee ee ee ee ee ee C. parryi var. nevadense Seeds <0.6 mm. in diameter; abaxial fruiting sepals usually ovate with a very narrow scarious margin or none; South Coast Ranges, California......... ACKNOWLEDGMENTS Our sincere thanks to Rebecca Bratcher and Lee M. Simpson for technical help and Mary Alice Kessler for help with georeferencing of some specimens. Funding for the study of the genus Calyptridium was provided by the Anza-Borrego Foundation Howie Weir Memorial Conservation Grant, the Frank Alverson and Mabel Meyers Memorial Scholarships, Jordan Dale Covin Memorial Travel Grant, the Sally Casanova California Pre-Doctoral Program, and the SDSU Field Stations Graduate Student Research Award. We also thank the following herbaria for allowing us to examine material: ARIZ, ASU, CAS-DS, JEPS, RSA, SD, SDSU, UC, UCR, and UTC LITERATURE CITED APG III. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG. III Botanical Journal of the Linnean Society 161:105—121. C. parryi var. hesseae CRONQUIST, A. 1978. Once again, what is a species? Pp. 3—20 in Ramberger J. A. (ed.), Biosystematics in agriculture. Allanheld & Osmun, Montclair, NJ. 1988. The evolution and classification of flowering plants. Ed. 2. New York Botanic Garden, New York, NY. GUILLIAMS, C. M. 2009. Phylogenetic reconstruction, character evolution, and conservation in the genus Calyptridium (Montiaceae) M.S. thesis. San Diego State University, San Diego, CA. RIEMANN, H. AND E. EZCURRA. 2007. Endemic regions of the vascular flora of the peninsula of Baja Califor- nia, Mexico. Journal of Vegetation Science 18:327— 330. Stmpson, M. G., M. SILVEIRA, AND C. M. GUILLIAMS. 2010. Taxonomy of Calyptridium parryi (Montia- ceae) Madrono 57:145—160. THORNE, R. F., R. V. MORAN, AND R. A. MINNICH. 2010. Vascular plants of the high Sierra San Pedro Martir, Baja California, Mexico: an annotated checklist. Aliso 28:1—50. MADRONO, Vol. 58, No. 4, pp. 267-272, 2011 NOMENCLATURAL KANKEDORTS IN PHACELIA (BORAGINACEAE: HYDROPHYLLOIDEAE) GENEVIEVE K. WALDEN! AND ROBERT PATTERSON Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, CA 94132 gkwalden@berkeley.edu ABSTRACT The following nomenclatural combinations in Phacelia (Boraginaceae) are established: P. campanularia A. Gray var. vasiformis (G. W. Gillett) Walden & R. Patterson, stat. nov.; P. imbricata Greene var. bernardina (Greene) Walden & R. Patterson, stat. nov.; P. imbricata Greene var. patula (Greene) Walden & R. Patterson, stat. nov.; and P. nemoralis Greene var. oregonensis (Heckard) Walden & R. Patterson, stat. nov. Typification and clarification of nomenclature is provided for Phacelia floribunda Greene and Phacelia phyllomanica A. Gray. Synonymies are provided for all names. Key Words: California flora, Channel Islands, Hydrophyllaceae, Isla Guadalupe, Oregon flora. Phacelia Juss. has been traditionally included within Hydrophyllaceae, with molecular studies placing it in Hydrophylloideae within the larger Boraginaceae (Burnett 1835; Ferguson 1998; APG II 2003). Within the history of Phacelia, divisions have received varied amounts of atten- tion resulting in an unwieldy system of nonstan- dard ranks (Walden 2010). Nomenclatural changes are proposed here in an effort to standardize infraspecific rank and in anticipation of the Flora of North America North of Mexico treatment of the genus. Taxa are presented in alphabetical order with keys, synonymies, and lectotypifications where necessary. HISTORY The divisions within Phacelia have benefited from, and have been complicated by, multiple researchers. Gray (1875a, 1875b, 1878, 1886) can be credited for the major impetus to assign divisional rank within the genus, which has only been comprehensively treated one time by Brand (1913). Gillett (1968) and Lee (1986) both used “subgroup” to refer to divisions within subg. Cosmanthus and species group Humiles respec- tively, but these “‘subgroups” are not equivalent and represent different syntheses of evolutionary relationships. The inclusion of both subspecies and varieties within the genus presents similar difficulties. Some taxonomists have been clear in their definition and application of the rank subspecies, such as Gillett (1955) in his treatments of sect. Whitlavia and sect. Gymnobythus, or Heckard (1960) in his treatment of the P. magellanica complex; other authors, like Brand (1913), applied various infraspecific ranks ‘Present address: Department of Integrative Biology, University of California, Berkeley, CA 94720 throughout the genus. These previous efforts provide rich resources of names to consider when forming regional floras, with the overall trend in the genus toward infraspecific rank of variety, with few taxa not having a valid published varietal name. Change of subspecies names to varietal names is a remedy toward standardiza- tion of infraspecific rank in Phacelia. PHACELIA CAMPANULARIA Phacelia campanularia A. Gray consists of diploid plants (7 = 11), and has been championed in horticulture for the large, blue corollas, but the profuse glandular hairs can cause contact derma- titis (Curtis 1884; Munz 1932; Reynolds and Rodriguez 1986). TAXONOMIC TREATMENT Phacelia campanularia A. Gray, Syn. Fl. N. Amer. 2. 1. 164. 1878. Phacelia minor Thell. var. campanularia (A. Gray) Jeps., Man. Fl. PI. Calif. 819. 1925.—Type: USA, California, Riverside Co., White Water, on the left bank of the Whitewater [River] and perhaps 2 miles from, its exit, 1876, C. °C. Parry and J. G. Lemmon 263 (lectotype: US 1338221, designat- ed by G. W. Gillett 1955; isosyntypes: F 153461!, F 314052!, GH 92313, K, MO 2518983-—2235139 digital image!, MO 2518982—2235140 digital image!, NY 83955!, NY 83956!, NY 83957, NY 83958!, PH 759569— 28225 digital image!, UC 335625)!). Phacelia parryi Torr. var. celata Jeps. & Hoover ex Jeps., Fl. Calif. 3: 276. 1943.—Type: USA, California, Riverside Co., Colorado Desert, Whitewater River, San Gorgonio Pass, abun- dant in coarse gravel, flowers purple, 1907, S. B. Parish s.n. (holotype: JEPS 2792!). 268 The other syntype of P. campanularia cited in Gray’s protologue was identified by Gillett (1955) as a specimen of hybrid Phacelia minor X P. parryi (USA, California, San Diego Co., [Ori- flamme Canyon], 1881, D. Cleveland s.n. [syntype GH; isosyntype F)). Phacelia campanularia A. Gray var. vasiformis (G. W. Gillett) Walden & R. Patt., stat. nov. Phacelia campanularia A. Gray subsp. vasifor- mis G. W. Gillett, Univ. Calif. Publ. Bot. 28: 64. 1955.—Type: USA, California, San Bernardino Co., near Bonanza [Bonanza King Mine], Providence Mtns., rocky hillside, 2500 ft, 30 Mar 1920, P. A. Munz and R. D. Harwood 3446 (holotype: POM 7702!; isotypes: DS 122640!, UC 219293!, US 1223578 digital image!). Key to Varieties of Phacelia campanularia 1. Leaf blades 20-70 mm; petioles 10-100 mm; cymes 5-20 cm; corollas campanulate to rotate, (6—-)15-30 mm; styles 20-35 mm, branched 1/3—1/2 length....... var. campanularia 1’. Leaf blades (20—)30—-100 mm; petioles (10—)50— 200 mm; cymes 10—50 cm; corollas funnelform to funnelform-campanulate, (15—)25—40 mm; styles 30-45 mm, branched 1/4-1/2 length. . . Fide helen eas ea ie, DE ate ane var. vasiformis Distribution Gillett (1955) noted that the subspecies of P. campanularia (sea level to 1600 m) were separated within the Salton Trough by the Imperial Valley and Coachella graben, a geographic limitation examined through additional collecting and molecular systematic studies (Wallace 1990; Hansen et al. 2009). PHACELIA IMBRICATA Phacelia imbricata Greene is variable and widespread, with diploids (n = 11) and tetra- ploids (n = 22) (Heckard 1960). Variation in pubescence is described in the key. TAXONOMIC TREATMENT Phacelia imbricata Greene, Erythea 1: 127. 1893. Phacelia californica Cham. var. imbricata (Greene) Jeps., Fl. W. Calif. 439. 1901.—Type: USA, California, Napa Co., near St. Helena, Jun 1891, FE. L. Greene s.n. (holotype: NDG 29239; isotype: UC 24385!; photographs of holotype UC 1055963!, UC 1471530). Phacelia circinata (Willd.) Jacq. var. calycosa A. Gray, Proc. Amer. Acad. Arts 10: 317. 1875. Phacelia californica Cham. var. calycosa (A. Gray) Dundas, Bull. S. Calif. Acad. 33: 158. 1935. Phacelia magellanica (Lam.) Coville var. calycosa (A. Gray) Jeps. & V. L. Bailey, FI. Calif. 3: 246. 1943.—Type: USA, California, MADRONO [Vol. 58 Mariposa Co., foothills to Yosemite, 1872, A. Gray s.n. (lectotype, designated by Dundas 1934: GH). Phacelia stimulans Eastw., Proc. Calif. Acad. Sci. ser. 3, Bot. 2: 291. 1902.—Type: USA, Cali- fornia, Fresno Co., south fork of King’s River, Kings River Canyon [not far from the swampy meadow near where campers stop on the way to Bubbs Creek], 1-13 Jul 1899, A. Eastwood s.n. (holotype: CAS 942!; photograph of holotype UC 657552!). Phacelia imbricata Greene var. condensata Brand, Univ. Calif. Publ. Bot. 4: 220. 1912.—Type: USA, California, Mariposa Co., Yosemite Valley [Die haufigste Form], n.d., H. Bolander s.n., (lectotype, here designated: G [Herb. Boissier]). Syntype: USA, California, Amador Co., New York Falls, 700 m, 3 May 1893, G. Hansen 1283 (B [likely destroyed]; isosyntypes: UC 192183!, JE 1635 digital image! [1500 ft, 1 May 1893)). Phacelia imbricata Greene subvar. hansenii Brand, Univ. Calif. Publ. Bot. 4: 220. 1912.—Type: USA, California, Amador Co., Irishtown, 500 m, 1 May 1893, G. Hansen 1283 (holotype: G [Herb. Boissier]). Phacelia imbricata Greene var. caudata Brand, Univ. Calif. Publ. Bot. 4: 220. 1912.—Type: USA, California, El Dorado Co., Natoma Ditch, 2 May 1909, K. Brandegee s.n. (holotype: UC 130944!). No specimens were cited in the protologue for P. circinata. var. calycosa, but four specimes were cited in the description published in 1876 (Gray 1875a; Brewer et al. 1876). All four specimens were collected prior to 1875 and deposited at GH, and although not cited in the protologue, make up the original material upon which the descrip- tion was based. In his revision of the group, Dundas (1934) suggested ’Gray’s own collection be designated as the type.” Jepson (1943) designated Torrey 343 (sic) as the lectotype, based on it being first cited by Gray in Bot. Calif. (Brewer et al. 1876). Torrey 343 may be an error for Torrey 348 (the label of Jepson 18922 (JEPS 2144) noted “Exact match for type, Borax Lake, Torrey 348, in Gray Herb — W.L.J.”’). The lectotypification by Dundas is recognized here as having priority over that by Jepson. Brand (1912) cited only Bolander s.n. and Hansen 1283 in the protologue of P. imbricata var. condensata, without designating a holotype. The types and other coilections of Phacelia deposited at the Berlin Herbarium were destroyed in 1943 (Hiepko 1978, 1987). A duplicate of Hansen 1283 was deposited at UC, and a specimen with an earlier collection date at JE. The specimen Hansen 1283 “in Herb. Bois- sier, nicht in Herb. Berlin’? was designated the type of P. imbricata subvar. hansenii (Brand 1912). This specimen likely represents a different 2011] gathering (locality Irishtown) than P. imbricata var. condensata (locality New York Falls) under the same collection number (Hansen 1283) (Brand 1912). Phacelia imbricata Greene var. bernardina (Greene) Walden & R. Patt., stat. nov. Phacelia virgata Greene var. (?) bernardina Greene, Erythea 4: 55. 1896. Phacelia magellanica (Lam.) Coville forma bernardina (Greene) Brand, Univ. Calif. Publ. Bot. 4: 217-218. 1912. Phacelia californica Cham. forma bernar- dina (Greene) J. F. Macbr., Contr. Gray Herb. 49: 36. 1917. Phacelia californica Cham. var. bernardina (Greene) Jeps., Man. Fl. Pl. Calif. 820. 1925. Phacelia imbricata Greene subsp. bernardina (Greene) Heckard, Univ. Calif. Publ. Bot. 32: 44. 1960.—Type: USA, Califor- nia, San Bernardino Co., San Bernardino, 1 Jun 1887, S. B. Parish and W. F. Parish s.n. (holotype: NDG 40515; photographs of holo- type UC: 1055967!, UC 1471528!). Phacelia imbricata Greene var. patula (Brand) Walden & R. Patt., stat. nov. Phacelia magella- nica (Lam.) Coville forma patula Brand, Univ. Calif. Publ. Bot. 4: 217, 219. 1912. Phacelia californica Cham. var. patula (Brand) Jeps., Man. FI. Pl. Calif. 820. 1925. Phacelia magella- nica (Lam.) Coville var. patula (Brand) Jeps., Fl. Calif. 3: 248. 1943. Phacelia imbricata Greene subsp. patu/a (Brand) Heckard, Univ. Calif. Publ. Bot. 32(1): 45. 1960.—Type: USA, California, San Diego Co., Stonewall Mine, Cuyamaca Mtns., 4600 ft, 5—7 Jun 1897, S. B. Parish 4423 [holotype: B [Herb. Brand], de- stroyed Mar 1943; lectotype designated by Heckard 1960: DS 135519!; isolectotypes: CAS 35175!, F 108608!, JEPS 2788!, MSC 68875!, UC 107578!, US 313417 digital image!). Phacelia oreopola Heckard, Univ. Calif. Publ. Bot. 32: 47, tab. 7a. 1960.—Type: USA, California, Los Angeles Co., San Gabriel Mountains, western side of Mt. Gleason, 1.5 miles below [Mt. Gleason] summit along road to Tujunga Canyon, 6000 ft, 15 June 1953, tetraploid n = 22, L. R. Heckard 505, with R. Bacigalupi and G. T. Robbins (holotype: UC 244089!; isotypes: DS 430574!, F 1515590!, JEPS 28420!, NY 83887!, RSA 134098!). Key to Varieties of Phacelia imbricata 1. Plants not mephitic, white-hispid to white- tomentose, rarely glandular; foliage gray-green to whitish; basal leaves pinnate with 1-3 pairs of leaflets and larger, entire terminal leaflet; corollas white or pale pink to lavender; imbricate calyx lobes lanceolate to oblong in AOL a ele ee var. patula 1’. Plants mephitic, hispid and glandular; foliage green to yellow-green to gray-green; basal leaves pinnate with 3-7 pairs of leaflets and larger, entire terminal leaflet; corollas white; imbricate calyx lobes ovate to obovate in fruit WALDEN AND PATTERSON: NOMENCLATURAL KANKEDORTS IN PHACELIA 269 2. Corolla lobes erect to incurved; inflores- cences loosely paniculate...... var. imbricata 2’. Corolla lobes erect to slightly divergent; inflorescences densely virgate . . var. bernardina Distribution Phacelia imbricata is distributed in the Sierra Nevada and Coast Ranges south to Baja California. Phacelia imbricata var. imbricata ranges from the Sierra Nevada to the Coast Ranges of California from 50—2500 m, generally occurring at elevations above P. imbricata var. bernardina in the San Bernardino and San Gabriel Mountains (500—2000 m). Phacelia im- bricata var. bernardina occurs in the San Bernar- dino, Santa Ana, and San Gabriel Mountains at 250-500 m. Phacelia imbricata var. patula ranges from the San Gabriel and San Bernardino Mountains (750-2500 m), to Sierra de Juarez and Sierra San Pedro Martir (725—1750 m). PHACELIA NEMORALIS Phacelia nemoralis Greene var. nemoralis con- sists of diploid plants (n = 11) and P. nemoralis var. oregonensis consists of tetraploid plants (n = 22) (Cave and Constance 1942, 1947; Heckard 1956). Phacelia nemoralis has both glandular hairs and a large and robust type of eglandular hairs, which Heckard termed “‘stinging’’ (Heckard 1960; Di Fulvio and Dottori 1995). TAXONOMIC TREATMENT Phacelia nemoralis Greene, Pittonia 1: 141. 1887. —Type: USA, California, Alameda Co., Oak- land Hills, 21 Jul 1887, E L. Greene s.n. (holotype: NDG 29300; photographs of holo- type UC 055960!, UC 1471533!). Phacelia nemoralis Greene var. oregonensis (Heckard) Walden & R. Patt., stat. nov. Phacelia nemoralis Greene subsp. oregonensis Heckard, Leafl. W. Bot. 8: 30. 1956.—Type: USA, Oregon, Multnomah Co., Sauvies Island, 4 miles north of Burlington ferry, n = 22, 28 May 1943, L. Constance and A. A. Beetle 2674 (holotype: UC 671710!; isotype: F 1331237!). Key to Varieties of Phacelia nemoralis 1. Stems 4—7 mm diam.; basal leaves pinnate with one pair of leaflets and larger, entire terminal leaflet; corollas green-white, 3.5—5 mm, limbs 3.5-4.5 mm diam.; seeds 1—3, 1.5-2 mm... . apse Fa, GAO ee a Gelato var. nemoralis 1’. Stems 7-10 mm diam.; basal leaves pinnate with 2-3 pairs of leaflets and larger, entire terminal leaflet; corollas yellow-white, 5—6 mm, limbs 4-5 mm diam.; seeds 1—2, 2—2.5 mm. . 2 oa, ithe, Saeco Ate estar Sere ee Oh ees var. oregonensis 2710 Distribution Phacelia nemoralis Greene var. nemoralis rang- es from San Luis Obispo Co. to Sonoma Co. (central California) from 50-700 m; P. nemoralis var. oregonensis ranges from northwestern Cali- fornia (Humboldt and Del Norte counties) through western Oregon to western Washington from sea level to 800 m (Heckard 1956, 1960). PHACELIA FLORIBUNDA AND PHACELIA PHYLLOMANICA Phacelia floribunda Greene and Phacelia phyl- lomanica A. Gray are island endemics and infrequently collected. Heteromorphic calyx lobes are common in Phacelia, although the shallow to deep pinnate lobing of calyx lobes readily distinguishes these two taxa in the field. TAXONOMIC TREATMENT Phacelia floribunda Greene, Bulletin of the California Academy of Sciences 1: 200. 1885.—Type: MEXICO, Baja California, Gua- dalupe Island, “‘part of type’, 20 April 1885, 7: S. Brandegee s.n. (lectotype, here designated: UC 107314!). Phacelia phyllomanica A. Gray var. interrupta A. Gray, Proc. Amer. Acad. Arts 11:87. 1876.— Type: MEXICO, Baja California, Guadalupe Island, frequent in warm nooks 1n rocky ravines in the middle and at the south end, February to May [Watson, p. 118], 1875, E. Palmer 72 (lectotype, designated by R. Moran 1996: GH 00303824!; isolectotypes: MO 217214-399521 digital image!, MO 217212-399522 digital image!, NY 1239401!, NY 1239402!). Greene did not explicitly reference his own collection in the protologue of Phacelia floribun- da, although inclusion of locality and explicit lack of an alternate collector indicated a personal collection. The frontispiece to the Bulletin of the California Academy of Sciences noted that all types of new species were deposited at CAS, which was prior to the 1906 earthquake (Greene 1885). Eastwood (1929) did not locate any types of P. floribunda or P. phyllomanica in CAS collections after the earthquake. Howell (1943) documented types of Phacelia described by Greene and deposited at NDG; he was unable to locate the type for P. floribunda. Jepson (1943) cited Greene as collector of the type, and although not definitive, this seemed to indicate that a type specimen was still in existence and possibly examined during his study at UC. A specimen at UC stamped as part of the Brande- gee Herbarium, has note “‘part of type’ (México: Baja California, Guadalupe Island, 20 April 1885, “part of type’, 7. S. Brandegee s.n. UC 107314!). This specimen was annotated by Brand MADRONO [Vol. 58 (‘“!Br’, handwritten on sheet), and cited in the P. floribunda treatment, along with Palmer 72 and A. W. Anthony s.n. (Brand 1913). Palmer’s specimen was cited in synonymy, and Anthony traveled to Guadalupe Island in March to June 1897, after publication of Greene’s protologue, and these collections do not make up original material for P. floribunda. Greene commonly sent fragments of specimens for determinations to the Brandegees, and when the Brandegee Herbarium was given to the University of California, Berkeley it was noted that duplicates and fragments of types were well represented (Bran- degee 1893; Allen 1907). The specimen label corresponds to the time period when Greene visited Guadalupe Island in April 1885, a trip that the Brandegees did not attend. The label date could be a transcription error. The specimen could be a later trip by the Brandegees to Guadalupe Island in March of 1897. Although without annotation by Greene or on a Greene label, this specimen likely represents a fragment of original material collected by Greene that informed his determination of P. floribunda as a distinct species from P. phyllomanica, sent to the Brandegees as a duplicate for their collection, with the presumed holotype specimen later destroyed in the CAS earthquake and fires. No specimens were cited in the protologue of P. phyllomanica var. interrupta, but Watson (1876) cited Palmer 72 and Brand (1913) cited Palmer 72 in synonymy, and Moran (1996) designated Palmer 72 as the holotype (GH). Moran’s lectotypification is correct for P. phyllomanica var. interrupta. Phacelia phyllomanica A. Gray, Proc. Amer. Acad. Arts 11:87. 1876.—Type: MEXICO, Baja California, Guadalupe Island, in large compact masses 1n the crevices of high rocks in the middle of the island, rare [Watson, p. 118], 1875, E. Palmer 71 (lectotype, here designated: GH 00093523!; isolectotypes: MO 217213-— 399521 digital image!, MO 217211—399524 digital image!, NY 83859!, NY 83860, NY 83861!, NY 83862!). No specimens were cited in the protologue, but Watson (1876) cited Palmer 71 as P. phylloma- nica, and Gray (1878) cited Palmer for P. phyllomanica as the original material which he based his description. Brand (1913) cited Palmer 71 and Franceschi 43 in his treatment of P. phyllomanica, collectors also cited by Eastwood (1929). Moran (1996) cited Palmer 72 as the type for P. phyllomanica, and his lectotypification is here corrected. Key to Phacelia floribunda and Phacelia phyllomanica 1. Annuals, (5—)10—60 cm; foliage dark green, sometimes appearing cinereous; inflorescenc- es hispid and densely glandular; corolla 2011] throats white to pale blue to lavender, lobes pale blue to lavender to purple, lobe margins spreading, not recurved or revolute; stamens included to slightly exsert, styles inclu- CG ae vance eerites teers ce ea Re Phacelia floribunda 1’. Subshrubs, 100—200 cm; foliage cinereous to canescent, sometimes dark green; inflorescenc- es densely hirsute and sparsely glandular; corolla throats and lobes blue-violet to purple, lobe margins sometimes revolute or recurved laterally; stamens exsert, styles exsert...... Ee ty Or arena ae oe ae eee Phacelia phyllomanica Distribution Phacelia floribunda is restricted in distribution to Guadalupe Island of Baja California and the Channel Islands of California. Phacelia floribun- da has been collected from Guadalupe Island, Outer Islet, and Islote Negro. In the Channel Islands the majority of collections are known from San Clemente Island and fewer collections from Santa Barbara Island. Phacelia phylloma- nica is known only from Guadalupe Island. ACKNOWLEDGMENTS We thank Nancy Morin, Bruce Baldwin, Dean Kelch, Beth Weil, Andrew Doran, Kim Kersh, Ana Penny, Deb Trock, Frank Cipriano, the Patterson and Baldwin graduate labs, Mei Griebenow, Jim Linnberg, and Trigger (service dog of GK W) for their support and patience during these studies. This paper represents a portion of a master’s thesis by GK W and is in support of the FNANM treatment of Phacelia by both authors. Curators at herbaria cited were kind and generous with access, loans, and information regarding collections, for which we are grateful. We are pleased to thank Brian Munson, Ann Marie Graham, Jim McKenzie, Emily Howe, Emma Havstad, Korie Merrill, Joel Shute, Steve Junak, and Jeanne Marie Acceturo for the opportunity to visit populations of Phacelia floribunda and the SCI herbarium during the Jepson Workshop on San Clemente Island (April 2011, GKW). Funding was generously provided in part by NSF GRFP and UC Berkeley Chancellor’s Fellowships to GKW. We appreciate the helpful comments by anonymous review- ers, the editors, and John Strother. LITERATURE CITED ALLEN, A. H. 1907. The Brandegee herbarium and library. The University of California Chronicle, an official record 9:73—76. APG II. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141:399-436. BRAND, A. 1912. Die Hydrophyllaceen der Sierra Nevada. University of California Publications in Botany 4:209—227. . 1913. Hydrophyllaceae. A. 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Sur les relations dans leur port ou leur aspect, que les plantes de certaines contrées ont entrélles, et sur une nouvelle espéce d’Hydrophylle. Hydrophyllum Magellanicum. Jour- nal dhistoire naturelle 1:371—376, plate 19. LEE, G. J. 1986. Systematic studies in the Phacelia humilis group (Hydrophyllaceae): Corolla venation patterns. American Journal of Botany 73:230—235. MORAN, R. 1996. The flora of Guadalupe Island, Mexico. Memoirs of the California Academy of Sciences 19:1—185. MADRONO [Vol. 58 Munz, P. A. 1932. Dermatitis produced by Phacelia (Hydrophyllaceae). Science 76:194. REYNOLDS, G. W. AND E. RODRIGUEZ. 1986. Derma- totoxic phenolics from glandular trichomes of Phacelia campanularia and P. pedicellata. Phyto- chemistry 25:1617—1619. WALDEN, G. K. 2010. Phylogeny of infrageneric relationships within Phacelia (Boraginaceae) in- ferred from chloroplast sequence data. M.S. thesis, San Francisco State University, San Francisco, CA. WALLACE, R. E. (ed.). 1990. The San Andreas fault system, California, US Geological Survey profes- sional paper 1515. US Government Printing Office, Washington, DC. WATSON, S. 1875. On the flora of Guadalupe Island, Lower California; list of a collection of plants from Guadalupe Island, made by Dr. Edward Palmer, with his notes upon them; descriptions of new species of plants, chiefly Californian, with revisions of certain genera. Proceedings of the American Academy of Arts and Sciences 11:105—148. YOUMANS, W. J. (ed.). 1894. Philibert Commerson, “‘the King’s naturalist’. Vol. 46 Popular Science Month- ly. D. Appleton and Company, New York, NY. MADRONO, Vol. 58, No. 4, pp. 273-274, 2011 REVIEW The Jepson Manual Vascular Plants of California, Second Edition. EDITED BY BRUCE G. BALDWIN, DOUGLAS H. GOLDMAN, DAVID J. KEIL, ROB ERT PATTERSON, THOMAS J. ROSATTI, AND DIETER H. WILKEN. 2012. University of Califor- nia Press, Berkeley, CA. 1600 pp. ISBN 9780520253124, $100.00, hardcover. It has been 19 years since the Jepson Herbarium at UC Berkeley produced the last flora of California, The Jepson Manual, Higher Plants of California (Hickman 1993). During that time, the way that taxonomists work has changed substantially, with phylogenetic analysis and the use of molecular sequence data becoming stan- dard procedure. It was time to bring the Jepson Manual up to current taxonomic standards, and the Jepson Herbarium (led by convening editor Bruce Baldwin) stepped up to the plate once again, publishing the second edition of The Jepson Manual in January, 2012. The book, available from the University of California press, is also available in electronic form, for those who want to use it on a Kindle or iPad. Three hundred and thirty two authors or editors are listed as contributing to the 1568- page, revised edition, which treats more than 6500 species, subspecies, and varieties. Some taxonomic treatments are unchanged from the 1993 edition, but many have been totally overhauled or at least changed enough to accommodate the approximately 310 additions to the California flora recognized in the manual. The book design is handsome, with a hard- back glossy cover, a handy ruler, and extra pages in the back for taking notes, web addresses and a geographic map inside the front cover, and a revamped list of conventions, including a useful section called *““Things to Remember When Using this Book.” Rare plants are now designated with a Star, invasives with a diamond, flowering times have been added (hurrah!), and _ horticultural information deleted (but is available online at the Jepson Herbarium website). The section on past climates and vegetation of California is completely rewritten by Constance Millar, and the geographic subdivisions section has been revised with a finessed map. If you open the back cover, you are greeted with a recent classification of the vascular plant families in the manual, including a phylogenetic tree. The plant families are divided into eight major groups: Lycophytes, Ferns, Gymnosperms, Nymphaeales, Magnoliids, Ceratophyllales, Eu- dicots, and Monocots. Within the body of the book, families, genera, and species are arranged alphabetically within these major groups, which differ greatly in size (from Ceratophyllales with 1 species to Eudicots with 4723). Part of the philosophy of the manual is to recognize monophyletic taxa as much as possible, which is why the 1993 separation of flowering plants into Dicotyledons and Monocotyledons had to be changed. The paraphyletic nature of the Dicots (at least in studies based on chloroplast DNA sequences) has resulted in their break-up into Nymphaeales, Magnoliids, Ceratophyllales, and Eudicots. Although the plant families and their page numbers are shown at the tips of the branches of the phylogenetic tree on the endpa- per, the listing is not alphabetical. A shortcut index, like that inside the cover of A California Flora and Supplement (Munz 1973), would be a useful feature for locating family treatments dispersed among the major groups. This more fine-grained monophyletic taxono- my is echoed throughout the manual; evidence has accumulated that some prominent genera from earlier treatments are paraphyletic or polyphyletic. Many of these have been divided into smaller monophyletic genera. Lotus, for example, has been split into three genera with the name Lotus now restricted to Old World species; native species are assigned to Acmispon and Hosackia, names resurrected from the 19th century. Camissonia has been divided into nine genera, Rhamnus into two, Gilia into five, Polygonum into five, and Potentilla into four. Linanthus has been split into Leptosiphon and Linanthus, but Leptodactylon is merged into Linanthus. The genus Aster has been broken up into seven genera, none of which is called Aster, while Gnaphalium has been broken up into four genera, Hemizonia into three, and Madia into six. There are a lot of new taxonomic concepts and names to learn. One of the curious problems with this monophyletic taxonomy in an _ alphabetically arranged book is that it is hard to figure out where a species has gone, since its new genus is not situated near its old genus and other close relatives. However, there is always the index. Unlike the 1993 manual, which had a “‘name change” section in the back of the book, the index serves that purpose in the new edition. Like the 1993 edition, not every species name is in the index (because the book is arranged alphabeti- cally), but you will find 1993 species names that have changed and the pages where they are listed in synonymy. As a space-saving measure, names already synonymized in the first edition are not included in the second, but the website for the 274 Index to California Plant Names is provided inside the front cover. Following a monophyletic philosophy did not always lead to the splitting of taxa in the new manual. The grass treatment by J. Travis Columbus and James P. Smith resulted in the lumping of Vul/pia and Lolium into Festuca and the lumping of all the needle grasses back into the genus Stipa. Many Californians are doing a happy dance now that their state grass is once again called Stipa pulchra. The downside of this monophyletic treatment of the grasses is that it no longer agrees with the Flora North America grass treatment, which was edited by Mary Barkworth and did not follow a monophyletic philosophy. Lumping and splitting and wholesale recon- figuration occurred in the circumscriptions of some flowering plant families in the new edition. Hydrophyllaceae and Lennoaceae have been lumped with Boraginaceae, Asclepiadaceae 1s merged into Apocynaceae, Sterculiaceae into Malvaceae, and Lemnaceae into Araceae; Acer- aceae and Hippocastanaceae have been lumped into the mostly tropical Sapindaceae. However Portulacaceae has been divided into two families, Liliaceae into 12 families, and the Scrophular- laceae, Orobanchaceae, and Plantaginaceae have been totally reconfigured, with Mimu/us now in the family Phrymaceae. These changes are in line with the flowering plant classification currently taught in up-to-date taxonomy classes and follows the Angiosperm Phylogeny Group clas- sification that is continually updated at the Missouri Botanical Garden website. The number of taxon changes (including family/genus assignments, specific epithet chang- es, and author changes) are substantial, leading a friend of mine to lament, “Since I no longer know the name of any plant in California, I have decided to give up botany and switch to the classification of bagels.’ However, the new genus segregates actually highlight how endemic and special our flora is. We now have more endemic genera, which is a good thing when trying to protect them. For those who need to translate an existing plant list from 1993 names to 2012 names, Dick Moe of the University and Jepson Herbaria has developed a handy tool called the Jepson Online Concordance available at The Jepson Online Interchange [of] California Floristics MADRONO [Vol. 58 (make sure to capitalize genus names and correct misspellings). The content of the revised edition is not a big surprise to most active California botanists. The editors of the new Manual have been posting submitted taxonomic treatments online for public review for at least the past three years. Nearly all of the treatments had been posted by last spring. Comments on the treatments were well-received and many changes were made. The online review copy of the manual was replaced by the new Jepson eFlora in November, 2011, allowing anyone to use the new manual without purchasing the hardcopy book. The online version has handy links to other resources of the Jepson Online Interchange including species distribution maps based on the Consor- tium of California Herbaria dataset. In addition, the eFlora treats more taxa than the hardcopy. Users will note that in some keys in the hardcopy version, a taxon name appears in brackets. These taxa are waifs (not considered fully naturalized), and their species description only appears in the online version. The plan for the future is to add newly described taxa, revised nomenclature, range extensions, and other discoveries to the eFlora so that they become available to the public in a timely fashion. This public service to the botan- ical community keeps the Jepson Herbarium at the center of the botanical wheel in California, even as university funding continues to ebb away. California botanists have no idea how spoiled they are — so many states lack this commitment from their herbaria and therefore lack any type of state flora. The second edition of the Jepson Manual is a monumental work, one that every California botanist will use, and we thank the editors, authors, and the Jepson Herbarium for a job well done. —ELLEN DEAN, Department of Plant Sciences, Univer- sity of California, Davis, CA 95616; eadean@ucdavis. edu. LITERATURE CITED HICKMAN, J. C. (ed.). 1993. The Jepson manual: vascular plants of California. University of Califor- nia Press, Berkeley, CA. Munz, P. A. AND D. D. KEck. 1973. A California flora and supplement. University of California Press, Berkeley, CA. MADRONO, Vol. 58, No. 4, pp. 275-276, 2011 NOTEWORTHY COLLECTIONS OREGON The following collections document range extensions in ‘““Urbanizing Flora of Portland, Oregon, 1806—2008” (Christy et al. 2009); for brevity: ““Urbanizing Flora’’. EUONYMUS EUROPAEUS L. (CELASTRACEAE).— Multnomah Co., Portland, openish, brushy area, mostly on the S side of Springwater Corridor ca. 0.15—0.22 mi E of S.E. 111th Ave., well established by 1998, possibly from the Leach Botanical Garden or other gardens in the area where known to be cultivated, with Athyrium filix-femina, Corylus sp., Crataegus douglasii var. suksdorfii, C. monogyna, Epilobium sp., Galium aparine, Holcus lanatus, Hypericum perforatum, Phalaris arundinacea, Polystichum munitum, Rubus armeniacus, Salix lasiandra, Spiraea douglasii; elev. 210 ft, 1S May 1998, Marttala 4708 (UC, WS); 28 June 2006, Marttala 4708a (OSC, UC, WTU); 30 September 2006, Marttala 4708b (UC,WS); 2 December 2006, Marttala 4708c (CAS, HPSU, NY, OSC, REED, WTU): 6 May 2007, Marttala 4708d (OSC); 16 May 2007, Marttala 4708e (NY, OSC, REED, WTU); 19 May 2007, Marttala 4708f (HPSU). N_ side of Springwater Corridor ca. 100 ft E of MPII, ca. 0.16 mi W of S.E. 122nd Ave., among Phalaris arundinacea 1n grassy, +open area with brush and small trees, with Crataegus monogyna, Galium aparine, Ilex aquifolium, Oemleria cerasiformis, Rubus armenia- cus, elev. 210 ft, 29 September 2007, Marttala 5503 (OSC, REED). Previous knowledge. Previously reported from seven- teen states east of the Mississippi River (USDA, NRCS 2010). Significance. The first report from Oregon and first report in a state west of the Mississippi River, a range extension of about 2500 km was made in “Urbanizing Flora’. In late May 2010 the Oregon Flora Project Atlas posted a Eugene collection (just east of Autzen Stadium) 11 May 2005, B. Newhouse 2005—2006 (OSC) (Oregon Flora Project 2010), so now there are populations of Euonymus europaeus in Portland and Eugene, separated by ca. 160 km. POLYCARPON TETRAPHYLLUM (L.) L. var. TETRAPH YLLUM (CAR YOPHYLLACEAE).—Multnomah Co., Port- land, N.E. Couch St. between N.E. 11th and 12th Avenues and on adjacent 11th and 12th Avenues, 24 July 2003, Marttala 5420 (CAS, GH, HPSU, NY); 11 September 2003, Marttala 5420a (OSC, REED); 12 November 2004, Marttala 5420b (UC, US); 17 May 2006, Marttala 5420c (MO, WS, WTU). Around the intersection of S.E. Sandy Blvd. and I1th Ave. and Ankeny St., 17 June 2009, Marttala 5554 (OSC); 17 June 2009, Marttala 5553 (WTU); 18 July 2008, Marttala 5534 (OSC); 18 July 2008, Marttala 5535A (UC); 18 July 2008, Marttala 5535B (REED). S.E. Yamhill St. between 6th and 7th Ave. and 7th Ave. N of Yamhill St., 18 July 2008, Marttala 5533 (HPSU, NY); 17 June 2009, Marttala 5533a (OSC). N side of S.E. Belmont St. between Martin Luther King, Jr. Blvd. and Grand Ave. along the E bound Morrison Bridge off ramp, 17 July 2006, Marttala 5467 (OSC). S.E. Salmon St. just E of 2nd Ave. and 2nd Ave. from Salmon St. S on the E side of 2nd Ave. to ca. 115 ft from the intersection, adjacent to the Pratt and Whitney Tile Building, 13 May 2008, Marttala 5529 (OSC, REED, WTU). Habitat invariably sidewalk crevices and seams of sidewalk and buildings or streets; usually open areas, almost never on N side of buildings; elev. ca. 40—110 ft; most often with Cardamine oligosperma, Cerastium glomeratum, Poa annua, Polyg- onum aviculare, Sagina procumbens, Sonchus oleraceus, and Spergularia rubra. Previous knowledge. Reported from SW Oregon near Gold Beach (6 collections), e.g., Curry Co., Gold Beach, 42°24’27"N, 124°25'14.16”"W, 20 August 1998, Richard R. Halse, s.n. (OSC), and known from the historical record in Portland (Lower Albina, 15 September 1902, E. P. Sheldon 10307, OSC), and Hood River (along the Columbia River, Hood River, 23 July 1880, L. F. Henderson, s.n., OSC) (Oregon Flora Project 2010). Although Polycarpon tetraphyllum is given as an annual, it invariably seems to survive through all but the hardest of our usually mild winters. Significance. When published in the ‘“Urbanizing Flora” the Portland sites were ca. 365 km from the then nearest known populations and separated from the earlier Portland collections by over 100 years. The recent postings of the Richard R. Halse 7550 Arch Cape (Clatsop Co., 45°48'30.96"N, 123°57'43.92”W, 16 Sep- tember 2008, OSC) and Nick Otting, Danna Lytjen 1106 (Lane Co., 43°55'6.24"N, 123°0'44.28”"W, 5 June 2005, OSC) collections (Oregon Flora Project 2010) start to fill in the distribution of this weedy species, separations of about 100 and 160 km from Portland. The Portland sites cluster along a nearly 1.5 km long corridor, in part following a major arterial, Sandy Blvd., suggesting an avenue of dissemination. POTENTILLA RECTA L. (ROSACEAE).—Clackamas Co., dirt banks, sloping grass-forb meadow, and weedy flats, Molalla, TSS R6E sec. 20, NW “% SE 1/16; elev. ca. 1040 ft (ca. 315 m), open area to partial shade of cottonwoods, with Populus balsamifera ssp. trichocarpa, bearded Jris, Daucus carota, Sonchus sp., grasses, Cirsium arvense, C. vulgatum, Taraxacum officinale, Epilobium brachycarpum, Hypochaeris radicata, Leu- canthemum vulgare, Rubus armeniacus, R. ursinus, Lepidium sp., Narvarretia sp., Juniperus sp., Acer sp., Buddleja sp., 26 July 2008, Marttala 5536 (HPSU, WS); 17 August 2008, Marttala 5536a (NY, OSC, REED, WTU). Previous knowledge. Present in all but three U.S. states and widespread but erratically distributed in Oregon (Oregon Flora Project 2010; USDA, NRCS 2010). The nearest documented site is 3 mi. south of Halsey, (Linn Co., 44°21'36"N, 123°8'24"W, 13 July 1978, Gaylee Goodrich 43, OSC) (Oregon Flora Project 2010). This is listed as a Class B Noxious Weed by Oregon Department of Agriculture and as a “B” designated weed/Quarantine according to USDA, NRCS (2010). Significance. A range extension of about 100 km. SAMBUCUS NIGRA L. ssp. NIGRA (CAPRIFOLIA- CAE).—Multnomah Co., Portland, Brookside Wildlife Area, ca. 50-100 ft E of S.E. 110th Drive and ca. 325 ft N of Brookside Drive, +flat, open, grassy, brushy area with scattered trees, E of patches of large rocks, with grasses, Abies grandis, Spiraea douglasti, Fraxinus 276 latifolia, Berberis aquifolium, Thuja plicata, Robinia pseudo-acacia, Solanum dulcamara, Acer circinatum, Galium aparine, Rubus armeniacus, Vicia sativa ssp. nigra, V. hirsuta, Geranium dissectum, 27 August 2007, Marttala 5495 (HPSU, UC, WS); 22 September 2007, Marttala 5495a (NY, OSC, REED, WTU); 7 June 2008, Marttala 5495b (NY, OSC, REED, WTU). Previous knowledge. Known from three states on the east coast — Connecticut, Pennsylvania and Virginia (USDA, NRCS 2010). Significance. A range extension of about 3700 km. These plants are evidently relicts of cultivation, but they are prolific seeders and weedy; Sambucus nigra ssp. nigra 1s expected to spread. (Hogen 2003). SAXIFRAGA TRIDACTYLITES L. (SAXIFRAGA- CEAE).—Mutnomah Co., Portland, fenced vacant lot bounded by S.E. Taylor and Salmon Streets, S.E. Water Avenue and Interstate 5 and adjacent stretch of S.E. Taylor just E of small parking lot adjacent to Interstate 5, moss covered areas of asphalt and _ concrete (sidewalk), occasionally in thin, gritty soil as along sidewalk, with Brachythecium albicans, Bryum argen- teum, Ceratodon purpureus, Didymodon vinealis, Grim- mia pulvinata, Pseudoscleropodium purum, Rosulabryum capillare, Scleropodium cespitans, Syntrichia_ ruralis, Vulpia sp., Bromus rigidus, Hordeum murinum, Poa annua, Draba verna, Cardamine oligosperma, Senecio vulgaris, Stellaria media, Veronica arvensis, Cerastium glomeratum, C. semidecandrium, Acer macrophyllum, Arabidopsis thalliana, Daucus carota, Epilobium sp.., Hypochaeris radicata, Plantago lanceolata, Sonchus oleraceus, Trifolium dubium, Veronica arvensis, elev. ca. 30 ft, 8 April 2008, Marttala 5514 (CAS, GH, HPSU, NY, OSC, REED, UC); 9 April 2008, Marttala S5l4a (US, WTU); 25 April 2008, Marttala 5514b (CAS, GH, HPSU, NY, OSC, REED, UC, US, WTU); 14 May 2008, Marttala 5514c (BH, BRIT, BRY, GZU, MO, PE, UBC, WS). The vacant lot in which this population occurs is used for storage and transient parking. Despite many years of visiting and botanizing this site, no Saxifraga tridactylites was seen until 2008. Previous knowledge. Previously known in North America only from British Columbia, from Texada Island and sites in southern Vancouver Island near and west of Victoria (A. Ceska, Ceska Geobotanical Consulting, Victoria, BC; M. Fairbarns, Aruncus Consulting, Victoria, BC, and F. Lomer, Univ. British Columbia Herbarium, Vancouver, BC, personal com- munications to J. Christy, Portland State Univ., forwarded to author). Significance. First record in the continental U.S., ca. 330 km from nearest site in British Columbia, Canada. MADRONO [Vol. 58 In Europe, this species is expanding its range (Reisch 2007). SOLANUM LYCOPERSICUM L. var. LYCOPERSI- CUM (SOLANACEAE).—Multnomah Co., Portland, rocky, E bank of Willamette River ca. 350 ft S of Hawthorne Bridge, with Chenopodium ambrosioides, Rubus armeniacus, Rumex sp., Vicia sp., elev. 10—15 ft, plants did not survive the hard winter and high water of 2008-2009, 26 November 2008, Marttala 5543 (OSC, REED); 4 December 2008, Marttala 5543a (WTU). Washington Co., West Slope, TOIN, ROIW, NE1/4 of NE 1/4 Sec. 12, elev. 590 ft, sprouting nearly every summer from previous year’s seeds in garden beds and compost piles, with Malva neglecta, Kickxia elatine, Taraxacum officinale, Sonchus oleraceus, Portulaca oleracea, 28 Aug 2009, John A. Christy 10059 (OSC). Previous knowledge. Previously reported from Ore- gon (INVADERS Database System 2008; USDA, NRCS 2010) from Portland based on Suksdorf 1900 material (WS0000138469). Significance. Modern reports, more than 100 years after first collected. Since tomatoes regularly self-seed in gardens, the rarity of naturalized specimens in Oregon is fairly surprising, especially since USDA, NRCS (2010) shows them in forty states. —VERNON M. MARTTALA, 10811 S.E. Schiller St., Portland, OR 97266-3459. romanzoffivm@earthlink.net. LITERATURE CITED CHRISTY, J. A., A. Kimpo, V. MARTTALA, P. K. GADDIS, AND N. L. CHRISTY. 2009. Urbanizing flora of Portland, Oregon, 1806-2008. Occasional Paper 3. Native Plant Society of Oregon, P.O. Box 902, Eugene, OR. HOGEN, S 2003. FLORA, a gardener’s encyclopedia. Volume 2, L-Z. Timber Press, Portland, OR. OREGON FLORA PROJECT. 2010. Oregon plant atlas. Oregon State University, Corvallis, OR. Website http://www.oregonflora.org/atlas.php [accessed 31 May 2010]. REISCH, C. 2007. Genetic structure of Saxifraga tridactylites (Saxifragaceae) from natural and man-made _ habitats. Conservation Genetics 8:893-902. INVADERS DATABASE SYSTEM. 2008. The University of Montana, Missoula, MT.Website http://invader. dbs.umt.edu/ [accessed 4 December 2008]. USDA, NRCS. 2010. The PLANTS Database. National Plant Data Team. Greensboro, NC. Website http:// plants.usda.gov [accessed 2 June 2010]. MADRONO, Vol. 58, No. 4, p. 278, 2011 PRESIDENT’S REPORT FOR VOLUME 58 Dear CBS member, 2011 has been an excellent year for the California Botanical Society, and our successes only suggest more to come. Let me take this opportunity to review some of our achievements and to thank those involved. We redesigned our website (www.calbotsoc.org) and improved the acces- sibility of the information for our membership. We welcome your suggestions for continuing to improve this interface. In addition, many changes have occurred with Madrojo this year. Tim Lowrey has finished his term as Corresponding Editor and Matt Ritter from Cal Poly, San Luis Obispo has stepped in to replace him. On behalf of the society, I offer my sincere thanks to Tim for his fine work with the journal. He and Copy Editor Richard Whitkus have been instrumental in the conversion to an online manuscript submission and tracking process. Another exciting event regarding Madrono was that our proposal to JSTOR (an online archiving system) to place all of our back issues online was accepted. Sometime in the near future you will be able to access all of the older volumes through their website. In February, Second Vice-President Marc Los Huertos did an excellent job of organizing the Society’s annual banquet, held at California State University, Monterey Bay. We were treated to a talk by Josh Tewksbury, from the University of Washington, who spoke about why chilies (Capsicum spp.) are hot. For 2012, we did not holding a separate banquet meeting, but instead sponsored the banquet speaker for the California Native Plant Society’s January Conservation Conference in San Diego. Peter Raven, Director Emeritus of the Missouri Botanical Garden, was our invited speaker, and he gave a wonderful talk reflecting on the history of Western North American botany and what the future may hold. Anna Larsen has replaced Heather Driscoll as Corresponding Secretary for the Council after Heather left for an out of state position. The Council has worked particularly hard for you this year and I would especially like to thank Staci Markos for her various efforts on behalf of the Society. Thanks go, too, to Heather Driscoll and Kim Kersh for their constant management, especially with Society membership. And, finally, special thanks to Dean Kelch and Tom Schweich for all of their financial work and representing us at conferences. All of our Council members have been critically important this year. Given that the California Botanical Society was established in 1913, this April 2012 represents the 99‘? birthday of the Society. To celebrate the centennial of our Society, we are planning to organize a meeting in April 2013. We are looking at 2012 as the initiation of a year of field trips sponsored by the Society, so check the website often to find out when and where these will be held. Our membership base is the foundation of the Society, and your support allows us to promote botanical research and education. Increasing our membership is always a priority, so please continue to encourage your colleagues to join us and to publish in Madrojo. This is especially true of our younger colleagues. As we move online we hope to be more attractive to the younger cohorts of botanists more accustomed to this format. This year, in addition to encouraging other botanists you know to join the Society, we are also hoping to hear from you about your ideas for the celebration of our 100" year. Let us know by emailing or writing to any member of the Council. We’re certain that you have some incredible ideas for the centennial celebration! Also, please consider providing a sponsoring member- ship or subscription to a foreign scientist or scientific institution to support botanical research in economically depressed, developing countries. For more information on making such a gift, please contact Corresponding Secretary Anna Larsen (secretary@calbotsoc.org). The Society also sincerely welcomes gifts or other contributions to our endowment. V. Thomas Parker December 2011 MADRONO, Vol. 58, No. 4, p. 279, 2011 EDITORS’ REPORT FOR VOLUME 58 We are pleased to report the publication of this volume of Madrono by the California Botanical Society (CBS) in 2011. The journal continues to evolve as we get closer to the centennial year in 2013. Starting with the next volume, the CBS will introduce a “Point-of-View” section which will allow contributors to provide input on recent articles or air ideas related to western botany. We are hoping this will be a lively and informative forum. The Madrono page on the CSB web site is an up-to-date gateway for contributors, provides the link to the online submission page, and links to the eighty- year index. We continue to see an increase in the number of submissions and better turn around with reviewers. The efforts of numerous individuals are critical to the continuing quality of the journal. Among these, our Noteworthy Collections editor, Dieter Wilken; Steve Timbrook who has long provided the volume Index and Table of Contents; Annielaurie Seifert at Allen Press; and the enthusiastic support of the CBS executive council. Finally, we are extremely grateful to our contributors who provide interesting and insightful manuscripts, and our volunteer reviewers who take time from their busy schedules to assess the quality of submitted work. This year we received 43 new manuscripts and 39 were accepted for publication. Several manuscripts were also carried over from the previous year. The current volume includes 20 articles (including Notes), five new taxa, five Noteworthy Collections, and two Book Reviews. We appreciate the mix of submitted manuscripts across the spectrum of botanical sciences and anticipate continued submissions of novel and exciting work. As Editors, we have enjoyed our interactions with contributors and reviewers this past year. In the coming year, we have a new Corresponding Editor, Matt Ritter who brings a fresh perspective and has already taken on many responsibilities with the journal. For the coming year we are focused with brining the journal up to date so that we start the Centennial year on time and with exciting contributions related to the Centennial celebration. Tim Lowrey Richard Whitkus December 2011 MADRONO, Vol. 58, No. 4, p. 280, 2011 REVIEWERS OF MADRONO MANUSCRIPTS 2011 Daniel Austin Jan Beyers Andrew Bower Leo Bruederle William Buck James Carolin Curtis Clark Mark Egger Naomi Fraga Janet Franklin Brian Geils Patrick Gonzalez Henk Greven Shana Gross Benjamin Hall Richard Halse Gert Hansen Roxanne Hastings Takeo Horiguchi Philip Jenkins Leigh Johnson Eugene Kozloff Aaron Liston Joao Mieria-Neto Nancy Morin Robert Naczi Guy Nesom Daniel Nickrent Gilberto Ocampo Daniel Pinero Dalmau Robert Preston Jon Rebman James Reveal John Sawyer Leila Shultz Michael Simpson Jay Sobel John Strother David Tank Marcia Waterway William Weber Stanley Welsh Tom Wendt Mike Williams MADRONO, Vol. 58, No. 4, pp. 281—282, 2011 INDEX TO VOLUME 58 Classified entries: major subjects, key words, and results; botanical names (new names are in boldface); geographical areas; reviews, commentaries. Incidental references to taxa (including most lists and tables) are not indexed separately. Species appearing in Noteworthy Collections are indexed under name, family, and state or country. Authors and titles are listed alphabetically by author in the Table of Contents to the volume. Abies lowiana and A. magnifica dwarf mistletoes, 101. Alpine flora of Sierra Nevada, CA, diversity and biogeography, 153. Anemone tuberosa, noteworthy collection from MEXI- CO, 205. Apiaceae (see Lilaeopsis) Arceuthobium abietinum, comparison on Abies lowiana and A. magnifica, 101. Arizona: Floristic patterns on late Tertiary lacustrine deposits in Sonoran Desert, 123. Noteworthy collections: Artemisia pygmaea, Bursera microphylla, Fuirena simplex, Pholistoma membrana- ceum, Pulicaria paludosa, 64; Purshia glandulosa, 65. Arctostaphylos hooveri, lectotypification, 256. Artemisia pygmaea, noteworthy collection from AZ, 64. Asteraceae: Carduus pycnocephalus, effects of eradication and restoration treatments on, 207; Helianthus para- doxus and H. annuus, comparison of Na,SO,4 and NaCl effects on growth of, 145; Pentachaeta lyonii, impact of invasive annual plants on, 69. Noteworthy collections: Artemisia pygmaea, Pulicaria paludosa, 64. Astragalus kelseyae, new species from the Wasatch Mts. of UT, 135. Aucuba (see Garrya) Bangiaceae (see Porphyra) Bird dispersal of seeds (see Olea) Boraginaceae (see Phacelia) Brassicaceae (see Draba) Bursera microphylla, noteworthy collection from AZ, 64. Burseraceae (see Bursera) Cactaceae (see Cylindropuntia) Calamagrostis, morphology and phytogeography of native spp. from British Columbia, CANADA, 214. California: Arceuthobium abietinum, comparison on Abies lowiana and A. magnifica, 101; Arctostaphylos hooveri, lectotypification, 256; Carduus pycnocephalus, effects of eradication and restoration treatments on, 207; Ceano- thus roderickii, edaphic ecology and genetics of the gabbro-endemic shrub, 1; Clarkia unguiculata, pollen siring success, 78; Cylindropuntia X fosbergi, study of hybrid origin, 106; Darlingtonia californica, pollination biology, 22; diversity and biogeography of Sierra Nevada alpine flora, 153; gabbro soils, plant distribu- tions on, 113; Lilaeopsis masonii, taxonomic status, 131i; Olea europaea, fruit size relation to seed dispersal by birds, 86; Papaver californicum and Stylomecon heterophylla, systematics, phylogeny and evolution of, 92; Pentachaeta lyonii, impact of invasive annual plants on, 69; Pinus balfouriana, population ecology and demography, 234. New taxa: Grimmia vaginulata, 190; Mentzelia mono- ensis, 57. (See also Phacelia) Noteworthy collections: Polemonium carneum, 66; Porphyra suborbiculata, 201; Sequoiadendron gigan- teum, 202. Calyptridium parryi var. martirense, new taxon from Baja California, MEXICO, 258. CANADA (see Calamagrostis) Carduus pycnocephalus, effects of eradication and resto- ration treatments on, 207. Ceanothus roderickii, edaphic ecology and genetics of the gabbro-endemic shrub, |. Chromosome counts: Mentzelia thompsonii, 51. Clarkia unguiculata, pollen siring success, 78. Colorado (see Draba) Compositae (see Asteraceae) Crawford, Daniel J., dedication of Vol. 58 to, 283. Cupressaceae (see Sequoiadendron) Cylindropuntia X fosbergi, study of hybrid origin, 106. Cyperaceae (see Fuirena) Darlingtonia californica, pollination biology, 22. Desert floristic patterns on late Tertiary lacustrine deposits, 123. Draba weberi, noteworthy collection from CO, 204. Elymus alaskanus and E. violaceus, morphometric anal- ysis of variation between, 32. Edaphic ecology, (see Ceanothus) Editor’s Report for Vol. 58, 279. Ericaceae (see Arctostaphylos) Fabaceae (see Astragalus) Floristic patterns on late Tertiary lacustrine deposits in Sonoran Desert, 123. Fuirena simplex, noteworthy collection from AZ, 64. Gabbro soils, plant distributions on, 113. Garrya, molecular phylogenetics, 249. Garryaceae (see Garrya) Gramineae (see Poaceae) Grimmia vaginulata, new sp. from CA central coast, 190. Grimmiaceae (see Grimmia) Helianthus paradoxus and H. annuus, comparison of Na»SO, and NaCl effects on growth of, 145. Hydrophyllaceae (see Phacelia and Pholistoma) Invasive plants: Carduus pycnocephalus, effects of eradica- tion and restoration treatments on, 207; Olea europaea, fruit size relation to seed dispersal by birds, 86; Pentachaeta lyonii, impact of invasive annual plants on, 69. Keys: Astragalus spp. similar to A. kelseyae, 188; Calamagrostis of British Columbia, CANADA, 226; Calyptridium parryi and related spp., 266; Mentzelia Sect. Trachyphytum in Mono Co., CA, 62; Phacelia campanularia vars., 268; P. floribunda and P. phyllo- manica, 270; P. imbricate vars., P. nemoralis vars., 269. 0) MADRONO Lacustrine deposits, late Tertiary, floristic patterns on, 123: Leguminosae (see Fabaceae) Lilaeopsis masonii, taxonomic status, 131. Loasaceae (see Mentzelia) MEXICO (see Anemone and Calyptridium) Mentzelia: M. thompsonii, chromosome counts and taxonomy, 50. New taxon: M. monoensis, 57. Montiaceae (see Calyptridium) Olea europaea, fruit size relation to seed dispersal by birds, 86. Oleaceae (see Olea) Onagraceae (see Clarkia) Papaver californicum, systematics, phylogeny and evolu- tion of, 92. Papaveraceae (see Papaver and Stylomecon) Pentachaeta lyonii, impact of invasive annual plants on, 69. Phacelia: New combs. P. campanularia var. vasiformis, P. imbricate var. bernardina, P. i. var. patula, P. nemoralis var. oregonensis 268. Pholistoma membranaceum, noteworthy collection from AZ, 64. Pinaceae (see Pinus) Pinus balfouriana, population ecology and demography, 234. Poaceae (see Calamagrostis and Elymus) Polemoniaceae (see Polemonium) [Vol. 58 Polemonium carneum, noteworthy collection from CA, 66. Pollination biology (see Clarkia and Darlingtonia) Porphyra suborbiculata, noteworthy collection from CA, 201. President’s Report for Vol. 58, 278. Pulicaria paludosa, noteworthy collection from AZ, 64. Purshia glandulosa, noteworthy collection from AZ, 65. Reviews: Introduction to Chaparral by Ronald D. Quinn and Sterling C. Keeley, 199; The Jepson Manual Vascular Plants of California, Second Edition, ed. by Bruce G. Baldwin, et al., 273. Rhamnaceae (see Ceanothus) Rosaceae (see Purshia) Salt tolerance (see Helianthus) Sarraceniaceae (see Darlingtonia) Sequoiadendron giganteum, noteworthy collection from CA; 202. Sierra Nevada, CA, diversity and biogeography of alpine flora, 153. Stylomecon heterophylla, evolution of, 92. systematics, phylogeny and Tavares, Isabelle I., 1921—2011, In memorial, 67. Utah (see Astragalus) Viscaceae (see Arceuthobium) MADRONO, Vol. 58, No. 4, pp. 283-284, 2011 DEDICATION DANIEL J. CRAWFORD The California Botanical Society dedicates this volume of Madrono to Daniel J. Crawford, who is renowned not only for his studies of Coreopsis in California, western North America, Mexico and elsewhere, but also is acclaimed highly as a pioneering molecular plant systematist and an influential botanical role model. Dan was born in Columbus Junction, Iowa in 1942 and spent his early life on a farm where he was intrigued by the interesting assortment of roadside and agricultural weeds growing in the vicinity of his rural home. A devoted Hawkeye, he formalized his botanical training by pursuing three successive degrees at the University of Iowa. He received a B.A. degree in General Science in 1964 and in just two years, completed an impressive M.S. degree (1966) focusing on the Umbelliferae of Iowa. Dan continued his graduate studies under the supervision of Thomas E. Melchert and completed a Ph.D. degree in botany within three years (1969), a remarkable feat given that his dissertation involved a complex analysis of Mexican Coreopsis, which incorporated cytological, morphological and newly emerging “chemosystematic’ approaches. Dan began his career in the right place at the right time. While at U. Iowa, he overlapped with David E. Giannasi who was studying the flavonoid systematics of the genus Dahlia and the two became lifelong friends, eventually co-authoring a number of important chemo- systematic papers. Their advisor T. E. Melchert, a recent graduate of the University of Texas, had studied the cytology and chemosystematics of the composite genus Thelesperma under the supervision of noted synanther- ologist Billie Lee Turner. Turner and his colleague Ralph E. Alston were well-known to Dan by the plethora of chemosystematic studies they published in such presti- gious journals as Nature, PNAS and Science. In 1966 Melchert arranged for Dan to meet with Alston and Turner during a collecting trip to Mexico. That meeting between the 24-year-old botanist from Iowa and the two researchers at Austin must have been remarkably inspiring. Motivated by his advisor who was grappling with the precise delimitation of various genera within the Coreopsidineae, Dan found himself in the midst of two legendary botanists whose cutting-edge molecular research provided a tantalizing solution to elusive systematic problems. Dan has pursued a molec- ular approach to his systematic research ever since. However, he never abandoned his Levis or casual observations of country roadside flowers, but simply added a lab coat and laboratory bench to his repertoire. Incidentally, the lab coat remained a hallmark through- out his career and as a student, I never saw him in the lab without it. Dan’s expertise immediately secured him a job as assistant professor at the University of Wyoming, Laramie in 1969. It was there that his prolific publication record began with his first paper appearing in 1969 on a new species of Coreopsis from Mexico; shortly afterward he published several additional papers from his disserta- tion, which dealt with the cytology, flavonoid chemistry and morphology of Mexican Coreopsis. He quickly was promoted to Associate Professor (1973) and served as acting Head of the Department of Botany at Wyoming in 1974 and 1976. While at Laramie, Dan was secretary of Phytochem- ical Section of the Botanical Society of America (1975— 1976) and a member of the education committee (1978). He also began a systematic study of Chenopodium, publishing more than a half dozen papers from 1973— 1977 on the cytology, flavonoid chemistry, morphology and seed protein profiles for several species. Occasion- ally, a paper would appear on a group uncharacteristic of Dan’s usual focus, such as his chemical and morpholog- ical studies of Populus acuminatus (1974) and Arceutho- bium (1979). Such papers disclose Dan’s keen scientific curiosity, which surely accounts for much of his success. His seed protein research logically led to allozyme analyses, which quickly propelled him to the forefront of research in plant evolutionary systematics where he rapidly gained recognition as a worldwide authority. Dan’s first allozyme paper (on Chenopodium) appeared in 1977, co-authored by his post-doc Hugh Wilson. By virtue of his novel and exceptionally perceptive work, Dan’s rapidly escalating reputation resulted in an invitation to The Ohio State University as a Visiting Associate Professorship, for which he took a leave of absence from Wyoming during the 1977-1978 academic year. However, in 1977, the botanist from Columbus Junction moved to Columbus, Ohio to join the faculty of The Ohio State University permanently, where he advanced to full professor in 1980. Dan spent an ultra- productive 20 years at OSU before retiring in 2000. 284 At the Ohio State University, Dan joined forces with another B. L. Turner graduate and emerging Compositae researcher, Tod F. Stuessy. Dan and Tod went on to spearhead incredibly insightful research on the evolution of insular species by their collaborative work focusing on the flora of the Juan Fernandez Islands. The two continue to collaborate and their joint research has yielded dozens of articles encompassing nearly a thousand citations. This body of work has provided acute insight into the evolution of numerous genera on these islands. Dan’s more recent work has involved the Canary Islands, where he has focused on Tolpis (Asteraceae) and the conservation of the flora. Certainly one of the most amazing aspects of Dan’s career has been his ability to keep pace with the newly emerging technical developments in plant systematics. He has established himself in every emerging field, transitioning smoothly from flavonoid chemosystematics to allozyme analyses, and ultimately to various macro- molecular (DNA-based) approaches. As a result, Dan’s work has included an impressive array of studies incorporating such diverse approaches as RFLP, DNA- sequencing, and RAPDs analyses. He is known for his many excellent review articles, which began in 1978 with ‘Flavonoid chemistry and angiosperm evolution.” In 1985 he wrote a review entitled “‘Electrophoretic data and plant speciation’, which presented a comprehensive synthesis of allozyme data and their interpretation. The article literally has been cited a hundred times. Within five years he published the book: Plant Molecular Systematics: Macromolecular approaches, which was highly influential in that still developing field. Although he has contributed a significant amount of taxonomic literature, Dan’s true interests lie in the elucidation of plant speciation. Much of that interest derived from his studies in Coreopsis of closely related species pairs such as C. basilis/C. wrightii, C. gigantealC. maritima, C. nuecensis/C. nuecensoides and varieties of C. cyclocarpa. Dan has been a strong proponent of plant evolutionary study and has done much to encourage others to pursue research in this area. In 2000, he wrote an article entitled: ‘Plant Macromolecular Systematics in the Past 50 Years: One View’, which urged a more thorough study of “‘evolutionary processes and specia- tion” using DNA data. Aside from Coreopsis and island plants, Dan has studied a wide variety of other species in conjunction with a broad list of collaborators including many of his students and post-docs encompassing an amazing assortment of genera including Antennaria, Bidens, Calamagrostis, Calliopsis, Coreocarpus, Coreopsis, Lemna, Mabrya, Monarda, Paeonia, Polygonella, Sonchus, Tetramolopium, Trifolium, Trillium, Vittadinia and Wolffiella as some examples. Dan has long been an advocate of collaborative research, believing that molecular studies are most meaningful when coupled with more “traditional” approaches, especially fieldwork. He has collected several thousand plant specimens during his career and the MADRONO [Vol. 58 accompanying photo (provided by T. F. Stuessy) should remind everyone that he is not simply a laboratory scientist by any means, despite the highly technical nature of his work. I think that this philosophy is one reason why he has been able to get along well with people working in just about any field of biology. He always has held a deep respect for the work that others have contributed to his own studies and he always has valued the incorporation of diverse data sources in his own research. Such is exemplified by his lengthy collaboration with G. J. Anderson (University of Connecticut), who provided the additional dimension of breeding system information to Dan’s work on the evolution of island plants. Dan’s career has been studded with accolades, which include the presidencies of ASPT (1988) and Botanical Society of America (1996). In 1997 he was given the Asa Gray award, which represents the most prestigious honor bestowed by ASPT. Appropriately, he was presented the Alston Award for best phytochemical paper at the 1983 Botanical Society meetings. He received the BSA merit award in 1999 and a Centennial Award in 2006. His California connections include memberships in both the California Botanical Society and California Native Plant Society as well as service on the Editonal Board of Madrono from 1976-1981. After retiring from Ohio State, Dan secured an adjunct faculty position at the University of Kansas. He continues to be extraordinarily productive and currently is credited with two books and over 200 research articles. He also is involved with ‘“PlantingScience.org’’, which helps high school students become familiar with science. I remember vividly my arrival at The Ohio State University in 1980. When I first saw Dan milling about in his lab coat he terrified me. His research was spectacular and the intensely cerebral level of his graduate courses stunned me as well. In many ways I’m sure that I felt much like he did in 1966, awestruck by an encounter with one of the greatest personalities in all of systematic botany. However, I soon learned that behind the serious facade of a seasoned researcher was the soul of an absolutely wonderful person, unrivaled for his compassion, thoughtfulness and inspiration. He also liked professional wrestling! Dan’s scholarship provided a contagious air of “friendly academic competition” among his grad students. By his example, we all worked feverishly on our projects, each one of us wanting to achieve the same level of success as scientists, and also hoping to gain his seal of approval. He was the very best advisor anyone could hope for. Billie Turner (now 87) recently conveyed to me these words: “‘Dan has always been one of my favorite academics and, what else, human being. A fine example for man kind.’ How can I say it any better? Donald H. Les Professor of Ecology & Evolutionary Biology University of Connecticut Storrs, CT MADRONO A WEST AMERICAN JOURNAL OF BOTANY VOLUME LVIII 2011 BOARD OF EDITORS Class of: 2011—JAMIE KNEITEL, California State University, Sacramento, CA KEVIN RICE, University of California, Davis, CA 2012—GRETCHEN LEBUHN, San Francisco State University, CA ROBERT PATTERSON, San Francisco State University, CA 2013—-ERIC ROALSON, Washington State University, WA KRISTINA SCHIERENBECK, California State University, Chico, CA 2014—BRANDON PRATT, California State University, Bakersfield, CA TOM WENDT, University of Texas, Austin, TX Corresponding Editor—TIMOTHY LOWREY Museum of Southwestern Biology MSC03 2020 University of New Mexico Albuquerque, NM 87131-0001 madrono@unm.edu AND Copy Editor—RICHARD WHITKUS Department of Biology Sonoma State University 1801 E. Cotati Avenue Rohnert Park, CA 94928-3609 whitkus@sonoma.edu Published quarterly by the California Botanical Society, Inc. Life Sciences Building, University of California, Berkeley 94720 Printed by Allen Press, Inc., Lawrence, KS 66044 MADRONO VOLUME 58 TABLE OF CONTENTS Alexander, Earl B., Gabbro soils and plant distributions on them esi‘ i! Anderson, John L., Further floristics on late Tertiary lacustrine deposits it in the southern Arizona deserts ___ Anderson, John L., Noteworthy collections from Arizona. eee Aslan, Clare E., and Marcel Rejmanek, Smaller Olea europaea fruits have more potential dispersal implications for olive invasiveness in California Baldwin, Bruce G. (see Kadereit, Joachim W.) Barnett, Mia (see Mayer, Michael S., et al.) Blaylock, Christopher, Review of Jntroduction to California Chaparral by Ronald D. Quinn and Sterling C. WS CGI ao ees ae eee ci oe eee eet oe ia esses see ie once Brigham, Christy A. (see Moroney, Jolene R.) Brokaw, Joshua M., and Larry Hufford, A new species of Mentzelia (Loasaceae) from Mono County, 221110) 6 1: ee re eae ene ee eS Sar ee ee eer ee Yo, NT STS SO coy SEM NONE ORE hf Brokaw, Joshua M., Michael D. Windham and Larry Hufford, Chromosome counts and taxonomy of Mentzelia thompsonii Up@aSACCAG) ce .23. oo sis ee 2 ee Burge, Dylan O., Molecular phylogenetics of Garrya (Garryaceae) ee en Rr ee ate eau Burge, Dylan O., and Paul S. Manos, Edaphic ecology and genetics of the gabbro- endemic shrub Ceanothus VORCTICKIL CRADIMACE AC) © o.oo. eto he 8h pes en NOt Dc eng a ah soa Shek ete Sees tank ee ea ee Corbin, Beth Lowe, A new species of Astragalus from the Wasatch Mountains of Utah - Crumb, Esa K. (see Fiedler, Peggy L.) Dean, Ellen, Review of The Jepson Manual, Vascular Plants of California, Second Edition, edited by Bruce G. Bal wit U ae oo cece eg se oe en ws sts Slee ee pe ee i wel Se ee een Fallscheer, Robin (see Stubbs, Rebecca) Fiedler, Peggy L., Esa K. Crumb and A. Kate Knox, Reconsideration of the taxonomic status of Mason’s Lilaeopsis — a state-protected rare species in California Gromova, Anastasia (see Mayer, Michael S., et al.) Guilliams, C. Matt, Michael G. Simpson and Jon P. Rebman, Calyptridium parryi var. martirense (Montiaceae), a new taxon endemic to the Sierra de San Pedro Martir, Baja California, Mexico ____ Harrison, Kristen, and Richard J. Hebda, A morphometric analysis of variation between Elymus alaskanus and Elymus violaceae (Poaceae):Implications for recognition of taxa Hasenstab-Lehman, Kristen (see Mayer, Michael S., et al.) Hebda, Richard J. (see Harrison, Kristen) Hebda, Richard J. (see also Marr, Kendrick L.) Hufford, Larry (see Brokaw, Joshua M., and Larry Hufford) Hufford, Larry (see also Brokaw, Joshua M., Michael D. Windham and Larry Hufford) Hughey, Jeffery R., Noteworthy collection from Canora... 3.5. ta'l on oon. rase tenga laGele toca can eaeatea teased Kadereit, Joachim W., and Bruce G. Baldwin, Systematics, phylogeny and evolution of Papaver californicum and Stylomecon heterophylla (Papaveraceae) _ Keeley, Jon (see McGinnis, Thomas) Keil, David J., Lectotypification of Arctostaphylos hooveri (Ericaceae) Kellman, Kenneth, Grimmia vaginulata (Bryopsida, Grimmiaceae), a new species from the central coast of California oe eee p nee Knox, A. Kate (see Fiedler, Peggy L.) Kuhn, Bernadette, Noteworthy collection from Colorado Lahmeyer, Sean (see Vanderplank, Sula) Les, Donald H., Dedication of Volume 58 to Daniel J. Crawford __ Lippit, Molly (see Mayer, Michael S., et al.) Lowry, Tim, and Richard Whitkus, Editors’ Report for Volume 58° 4... 220. ).2.5 cit fesse eek Maloney, Patricia E., Population ecology and demography of an endemic subalpine conifer (Pinus balfouriana) with a disjunct distribution in California Manos, Paul S. (see Burge, Dylan O.) Marr, Kendrick L., Richard J. Hebda and Elizabeth Anne Zamluk, Morphological analysis and phytogeography of native Calamagrostis (Poaceae) from British Columbia, Canada and adjacent regions Marttala, Vernon M., Noteworthy collections from Oregon Mathiasen, Robert L., Morphological comparisons of white fir dwarf mistletoes in the Sierra Nevada and southern Cascade Mountains ee Mayer, Michael S., et al., Is Cylindropuntia X fosbergii (Cactaceae) ¢ a hybrid? a: ae McGinnis, Thomas and Jon Keeley, Effects of eradication and restoration treatments on Italian thistle COALQUUS DV CHOCCINOIISY a cctap icy race Boi RatGaaieig a ean alee ul een ane ae se ee, Meindl, George A., and Michael R. Mesler, Pollination biology of Darlingtonia californica (Sarraceniaceae), the California pitcher plant aed en Bese eee 199 213 131 258 32 201 92 2011] TABLE OF CONTENTS Mendez, M. O., and O. W. Van Auken, A comparison of the effects of NasSO, and NaCl on the growth of Helianthus paradoxus and Helianthus annuus (Asteraceae) Mesler, Michael R. (see Meindl, George A.) Moe, Richard L., In Memorial: Isabelle I. Tavares 1921-2011] : Moroney, Jolene R., Paula M. Schiffman and Christy A. Brigham, Invasive European annual plants impact a rare endemic sunflower . Parker, V. Thomas, President’s Report for Volume 58 Patterson, Robert (see Walden, Genevieve K.) Rebman, Jon P. (see Guilliams, C. Matt) Rebman, Jon P. (see also Mayer, Michael S., et al.) Rejymanek, Marcel (see Aslan, Clare E.) Rundel, Philip W., The diversity and biogeography of the alpine flora of the Sierra Nevada, California Schiffman, Paula M. (see Moroney, Jolene R.) Schmid, Mena (see Schmid, Rudolf) Schmid, Rudolf, and Mena Schmid, Noteworthy collection from California Simpson, Michael G. (see Guilliams, C. Matt) Smith-Heurta, Nancy L., and Frank C. Vasek, Pollen siring success in the California wildflower Clarkia unguiculata (Onagraceae) sere eaten siete eed uee Beatie te an sea ee Stubbs, Rebecca, and Robin Fallscheer, Noteworthy collection from California Van Auken, O.W. (see Mendez, M. O.) Vanderplank, Sula, and Sean Lahmeyer, Noteworthy collection from Mexico sss Vasek, Frank C. (see Smith-Heurta, Nancy L.) Walden, Genevieve K., and Robert Patterson, Nomenclatural kankedorts in Phacelia (Boraginaceae: Hydrophylloideae) == Be ee eee eee eee eee Whitkus, Richard (see Lowry, Tim) Windham, Michael D. (see Brokaw, Joshua M., Michael D. Windham and Larry Hufford) Zamluk, Elizabeth Anne (see Marr, Kendrick L.) ill DATES OF PUBLICATION OF MADRONO, VOLUME 58 Number |, pages 1—68, published 31 August 2011 Number 2, pages 69-130, published 13 January 2012 Number 3, pages 131—206, published 28 March 2012 Number 4, pages 207-284, published 3 July 2012 ee : 1 "=~ rah - Aw" . sae a SUBSCRIPTIONS — MEMBERSHIP The California Botanical Society has several membership types (individuals ($35 per year; family $40 per year; emeritus $27 per year; students $27 per year for a maximum of 7 years). Late fees may be assessed. Beginning in 2011, rates will increase by $5 for all membership types except life memberships, for which rates will increase by $100, and student memberships, which will not show a rate increase. Members of the Society receive MADRONO free. Institutional subscriptions to MADRONO are available ($70). Membership is based on a calendar year only. Life memberships are $750. 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