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Plant Communities and Floristic Diversity of the Emerald LakeBasin, Sequoia National Park, CaliforniaAuthor(s): Philip W. Rundel, Michael Neuman, and Peter RabenoldSource: Madroño, 56(3):184-198. 2009.Published By: California Botanical SocietyDOI: http://dx.doi.org/10.3120/0024-9637-56.3.184URL: http://www.bioone.org/doi/full/10.3120/0024-9637-56.3.184
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PLANT COMMUNITIES AND FLORISTIC DIVERSITY OF THE EMERALD LAKEBASIN, SEQUOIA NATIONAL PARK, CALIFORNIA
PHILIP W. RUNDEL, MICHAEL NEUMAN, AND PETER RABENOLD
Department of Ecology and Evolutionary Biology, University of California,Los Angeles CA 90095
ABSTRACT
The Emerald Lake Basin forms a subalpine watershed in the upper drainage of the Marble Fork ofthe Kaweah River in Sequoia National Park. The basin is 120 ha in area and covers an elevationalrange from 2804 m at Emerald Lake to 3416 m at the summit of Alta Peak. The flora of the basinincludes 202 vascular plant species, distributed into 132 genera and 41 families, with the Asteraceae(25 species) and Poaceae (23 species) as the largest families. Herbaceous perennials make up aboutthree-fourths of the flora, but unlike alpine habitats many of these are relatively tall upright species,particularly in wet habitats. The woody plant flora includes five tree species, all conifers, with Pinusmonticola as the dominant. There are 19 species of woody shrubs present, including six evergreen and13 winter deciduous species. Eight plant habitat types were delineated on the basis of geomorphicposition related to soil conditions and water availability that influence species composition (Billings1974). These were subalpine conifer forest, willow thicket, wet meadows, moist rock crevices, drymeadows, dry rock crevices, colluvium, and fellfields.
Key Words: Fellfield, Pinus jeffreyi, Sierra Nevada, subalpine flora, subalpine vegetation.
Subalpine and alpine basins of the SierraNevada provide important ecosystems not onlyfor their biodiversity but also for monitoringenvironmental impacts of global change resultingfrom higher temperatures, altered patterns ofsnowmelt, and the deposition of anthropogenicnitrogen and phosphorus. These high mountainsystems are sensitive to small changes in growingseason conditions of temperature and wateravailability (Bowman and Saestedt 2001; Malan-son et al. 2007). Additionally, there is littlepotential for nitrogen uptake by the limitedvegetation cover growing on shallow graniticsoils and the short growing season. Low levels ofnitrogen sequestration and deposition of anthro-pogenic nitrogen onto the winter snowpackproduces pulses of nitrogen associated withsnowmelt into aquatic systems, and thus an earlywarning indicator of nitrogen saturation fordownstream forested basins (Sickman et al.2003b; Williams et al. 1995). The Emerald LakeBasin in Sequoia National Park provides anexcellent case study for ecosystem stability in asmall subalpine watershed. The Emerald LakeWatershed Study (1984–1989) was organized bythe California Air Resources Board as a means ofbetter understanding the influence of atmosphericinputs of nutrients to basin processes andecosystem structure (Tonnessen 1991). Thesestudies have provided detailed and in many caseslong-term continuing databases to understandhydrologic flow and water balance (Kattelmannand Elder 1991), nutrient enrichment (Sickman etal. 2003a, b), solute chemistry of snowmelt andrunoff (Williams and Melack 1991), nitrogen
fluxes and transformations (Williams et al. 1995),and long-term models of hydrochemical respons-es (Wolford et al. 1996; Meixner et al. 2004). Ourobjective in this paper is to describe the floristicdiversity of the Emerald Lake Basin and tocharacterize the mesotopographic distribution ofplant communities within the basin.
MATERIALS AND METHODS
Study Site
The Emerald Lake watershed comprises arugged basin about 120 ha in area in the upperdrainage of the Marble Fork of the KaweahRiver above Tokopah Falls in Sequoia NationalPark (36u359490N lat., 118u409300W long.). Thebasin is roughly triangular in shape trendingnorthwest to southeast, with elevations rangingfrom 2804 m at Emerald Lake at the lower end ofthe basin to the apex of the triangle at the summitof Alta Peak at 3416 m (Figure 1). Two spursemanating from Alta Peak form ridges along thesouthern and northeastern boundaries of thedrainage, while the lower drainage boundaries areopen to the northwest.
The Emerald Lake Basin is a glacial cirque,carved from granitic parent material. Bedrockexposed by glacial scouring and frost actioncovers nearly half of the basin surface, with theremaining half covered by talus and thin soils inapproximately equal proportions (Sisson andMoore 1984; Tonnessen 1991). Exposed rockfaces in the basin contain many smaller fracturejoints, providing cracks where residual soils can
MADRONO, Vol. 56, No. 3, pp. 184–198, 2009
collect. The basin is typical of many subalpineand alpine lake basins in the Sierra Nevada withits weakly buffered calcium bicarbonate surfacewaters (Williams and Melack 1991).
Median slope angles in the basin are about 30u,but many of the upper slopes are considerablysteeper. A major fracture in the granite forms alarge joint that extends across the lower eleva-tions of the basin in a southwest to northeastdirection. Emerald Lake itself, 2.7 ha in area, is atarn formed by glacial quarrying of fracturedrock along this joint. There is a smaller lake at2963 m elevation above Emerald Lake near thegeographical center of the basin.
The climate of the Emerald Lake Basin istypical of the Mediterranean-type regime of thesouthern Sierra Nevada, with 75–90% the annualprecipitation falling as snow in the winter months(Stephenson 1990). For Emerald Lake, rainfallcomprises only about 10% of annual precipita-tion and occurs predominantly in the autumn.There are limited long-term data on precipitationlevels present in the basin. Mean annual precip-itation is about 1600 mm, but amounts are highlyvariable between years (Sickman et al. 2003a).Emerald Lake is typically covered by ice fromNovember to June or July. Snowmelt typicallybegins in April with peak discharge typicallyoccurring in June (Sickman et al. 2003b).Summers are generally dry except for an occa-sional convective storm associated with mon-soonal air masses from the east.
Field Studies
The field studies collecting the data presentedin this paper were carried out between June and
September in 1984, 1985, and 1987. Over thisperiod all parts of the basin were repeatedlyvisited to provide as complete an assessment aspossible of the vascular plant flora. These fieldstudies identified, characterized and mapped aseries of plant communities determined by acombination of their physiographic positionwithin the watershed, soil accumulation, andseasonal water availability. These communitiesbroadly resemble the mesotopographic gradientof alpine vegetation described by Billings (1974).Each of these communities had distinctivefloristic and plant life-form dominance. Noattempt was made to assign these communitiesto previous classifications of subalpine and alpinevegetation alliances and associations for theSierra Nevada (see Sawyer and Keeler-Wolf2007) as these applied relatively poorly to theEmerald Lake Basin because of broad ecotonalgradients between communities. Representativebiomass samples were collected from eachcommunity and a complete census and diametermeasurement of all trees in the basin wascompleted.
The scientific names used in this paper followthose of The Jepson Manual (Hickman 1993),with the exception of the familial classificationwhere the Liliaceae and Amaryllidaceae aretreated as separate families here.
RESULTS
Flora
The flora of the Emerald Lake Basin includes202 vascular plant species, distributed into 130genera and 42 families (Table 1). The largest
FIG. 1. The Emerald Lake Basin, Sequoia National Park, California. Emerald Lake at the lower margin of thebasin lies at 2804 m elevation while Alta peak at the southeast corner is the high point at 3416 m. Map by T.Meixler and used with permission.
2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 185
family present is the Asteraceae with 25 species,followed in order by the Poaceae (23 species),Scrophulariaceae (16 species), Cyperaceae (12species), and Rosaceae (11 species). These fivefamilies comprise 44% of the total flora. Notableis the absence of any species of Fabaceae, as otheralpine floras typically have this family wellrepresented (Rundel et al. 2008). At the genericlevel, Carex forms the largest group with 10species. Other genera with four or more speciesinclude Mimulus (6 species), Epilobium (4 spe-cies), and Saxifraga (4 species).
The Pteridophyta are represented by 11 speciesin the basin, all herbaceous perennials (Table 1,Appendix 1). The ferns are most important in wetmeadows and mesic rock crevices where Crypto-gamma acrostichoides, Athyrium alpestre, Cystop-teris fragilis, and Woodsia scopulorum are allcommon. Other pteridopytes are more typical ofdry meadows and dry rock crevices, as withAspidotis densa, Pellaea bridgesii, and Selaginellawatsonii. Ferns also are common in colluvialhabitats, as described below.
Five species of coniferous trees are present inthe basin (Table 1, Appendix 1), with four ofthese pines. More than 70% of these trees arePinus monticola, with Pinus contorta subsp.murrayana and P. balfouriana ssp. austrinacomprising 17% and 9.5% of the trees respective-ly. Only a few individuals of Pinus jeffreyi andAbies magnifica var. shastensis are present.
Angiosperms form more than 92% of the basinflora, with a total of 186 species (Table 1). Dicotsinclude three-fourths of the angiosperm total (139species), while monocots provide the otherquarter (47 species). Among the latter, graminoidlife forms of grasses, sedges, and rushes form thedominant element.
Life Forms
Woody plants make up only a moderateproportion of the flora of the Emerald LakeBasin. As described above, there are five speciesof evergreen trees present, all conifers. There are19 species of woody shrubs present, 9% of theflora, with six of these evergreen shrubs and theother 13 species as winter deciduous shrubs(Appendix 1). The evergreen shrubs include five
species of Ericaceae and Fagaceae—Arctostaph-ylos nevadensis, Chrysolepis sempervirens, Kalmiapolifolia, Ledum glandulosum, and Phyllodocebreweri. The first two of these are largelyrestricted to growing within the open pine standsof the basin. Kalmia polifolia is found in willowthickets and wet meadows, while L. glandulosumand P. breweri can be found in a variety ofcommunities. The sixth species of woody shrub,barely qualifying beyond being a subshrub, isEriogonum wrightii (Polygonaceae), a low-grow-ing dry meadow species.
The most prominent deciduous shrub in thebasin is Salix orestera, which forms extensivewillow thickets. A number of other deciduousshrubs are less abundant and generally exhibit alower stature. These include five species ofRosaceae—Amelanchier utahensis, Holodiscus mi-crophyllus, Prunus emarginata, Sorbus californica,and Spiraea densiflora. Also present are Loniceraconjugialis, L. involucrata, and Sambucus race-mosa (Caprifoliaceae), Ribes cereum and R.montigenum (Grossulariaceae), Jamesia america-na (Philadelphaceae), and Acer glabrum (Acer-aceae).
An additional eight species (4% of the flora) inthe basin can be classified as suffrutescentsubshrubs. These are Aster breweri and Erica-meria discoidea (Asteraceae), Epilobium canum(Onagraceae), Leptodactylon pungens, and Phloxdiffusa (Polemoniaceae), Primula suffrutescens(Primulaceae), and Penstemon newberryi (Scro-phulariaceae). These subshrubs are most charac-teristic of more xeric habitats such as pine forests,dry meadows, dry rock crevices and colluvialsites.
The overwhelming life forms for floristicdominance in the basin are herbaceous perenni-als, which comprise three-fourths of the totalflora. These are species that characteristically dieback to ground level at the end of each growingseason. Within this broadly defined category arerosettes, broad-leaved tussocks, perennial grami-noids, geophytes, mats and cushions, and bien-nials. The 156 species of herbaceous perennialsinclude 11 pteridophytes, seven monocot geo-phytes, 39 graminoids, and 98 herbaceous peren-nial dicots. At the subalpine elevations of theEmerald Lake Basin, many perennial dicotspecies are upright and relatively tall, in contrastto low-growing species that characterize thealpine environments.
The seven species of geophytes present are inthe Amaryllidaceae and Liliaceae (Allium cf.campanulatum, A. obtusum, Fritillara pinetorum,Lilium kelleyanum, Smilacina racemosa, Veratrumcalifornicum, Zygadenus venenosus). Six species ofherbaceous perennials are hemi-parasites. Theseare Castilleja applegatei, C. miniata, C. nana,Pedicularis atollens, and P. semibarbata (Scro-phulariaceae), and Orobanche uniflora (Oroban-
TABLE 1. SUMMARY OF THE FLORA OF THE EMERALD
LAKE BASIN, SEQUOIA NATIONAL PARK, SIERRA
NEVADA, CALIFORNIA.
Division Families Genera Species
Pteridophyta 4 10 11Coniferophyta 1 2 5Dicotyledonae 32 93 139Monocotyledonae 5 25 47Total 42 130 202
186 MADRONO [Vol. 56
chaceae). Pterospora andromedea (Ericaceae) is a‘‘saprophyte’’, epiparasitic through fungal myce-lium on vascular plant roots.
Annual plants are infrequent in the basin, with14 species comprising 7% of the flora. Annualspresent at Emerald Lake are Gnaphalium palustre(Asteraceae), Cryptantha sp. (Boraginaceae),Cuscuta californica (Cuscutaceae), Phacelia eise-nii (Hydrophyllaceae), Gayophyton humile (Ona-graceae), Gilia capillaris and Linanthus ciliatus(Polemoniaceae), Collinsia torreyi, Mimulus bre-weri, M. laciniatus, and M. whitneyi (Scrophular-iaceae), Galium bifolium (Rubiaceae), Saxifragabryophora (Saxifragaceae), and Muhlenbergiafiliformis (Poaceae). Cuscuta californica is anannual parasitic vine. The annual flora is mostapparent in dry meadow habitats, although thereare certainly conspicuous species present in bothwet meadows and wet rock crevices.
Vascular Plant Communities
Subalpine conifer forest. Although scatteredconifers, particularly pines, are widespread in theEmerald Lake Basin, an open community ofsubalpine conifer forest is well developed only ina small area in the extreme northeast corner ofthe basin near Emerald Lake. The four conifersspecies present in this community are Pinusjeffreyi. Abies magnifica, Pinus monticola, and afew individuals of Pinus contorta subsp. mur-rayana. The thin soils of the pine sites are sandyin texture and very poor in organic matter,allowing these soils to dry quickly in the summerwhen snowmelt ceases. This condition mayaccount for the relatively small number ofspecies, 30 in total, encountered in this commu-nity.
Beyond this small area of conifer forest,scattered individuals of Pinus monticola arewidespread on the mesic benches of the south-west-facing slopes of the basin. In addition tothese southwest-facing slopes, P. monticola alsooccurs as scattered trees on relatively flat drymeadows below the cirque of Alta Peak. Thereare several areas of the basin where P. monticolaappears to be slowly expanding its range bycolonizing dry meadows and dry crevice habitats.A fifth conifer, Pinus balfouriana subsp. austrina,is restricted to a relatively small area on the ridgerunning along the south- and southwest-facingmargin of the basin.
The understory of the conifer forest stands istypically dominated by Chrysolepis sempervirens,which locally exhibits ground cover up to 80–100%. In more open forest stands, there is a lowcover of dry meadow herbs, most notably Ivesiasantalinoides and Achillea millefolium. On thelower slopes of the basin, the stands of C.sempervirens mix with Phyllodoce breweri at themargins of dry meadow habitats. Also present in
these stands are Arctostaphylos nevadensis, Ame-lanchier utahensis, Prunus emarginata, and Sorbuscalifornica. Although Lonicera conjugialis may bepresent, it is more common growing along wetrock crevices. None of the conifer forest commu-nities has a unique herbaceous flora, with thosespecies present also typical of dry meadowhabitats.
Willow thicket. Willow thickets are widespreadacross the Emerald Lake Basin and dominatesignificant areas of the basin. Along with Salixorestera, the indicator species, there are acharacteristic set of understory shrubs and herbs,which typically cover 100% of the groundsurface. Characteristic species include ericaceousshrubs such as Ledum glandulosum, Kalmiapolifolia, Phyllodoce breweri, and Vacciniumnivictum. Tall herbs present include Seneciotriangularis, Epilobium angustifolium, Lilium kel-leyanum, Ligusticum grayi, and Calamagrostiscanadensis. The high productivity of leaf litter inthis habitat has lead to the development of thickand hydric organic soils. The willow thicketcommunity typically intergrades to wet meadowcommunities around its margins. During ourstudy we observed that willows appeared to becolonizing wet meadow and wet rock crevicehabitats in several areas. With a dominance ofwoody plant cover and associated tall herbaceousperennials, the species richness of willow thicketsis relatively low. Only 28 species were encoun-tered.
Wet meadows and moist rock crevices. Wetmeadow and moist rock crevice communitiesform diverse habitats in the basin, and exhibit anumber of distinctive associations. These associ-ations are typically distributed in relatively flatareas on west-facing slopes above Emerald Lakewhere soils accumulate to moderate depths andremain moist for much or all of the growingseason. Variations in soil texture, soil organicmatter content, and soil moisture dynamicsappear to be the primary physical factorsseparating different wet meadow associations.The largest areas of wet meadow habitat arelocated high in the basin, adjacent to and aboveParson’s Pond, where snowmelt keeps the soilssaturated well into the summer. Other large areasof wet meadows occur on a bench runningsouthwest from the major area of Pinus mon-ticola, and adjacent to willow thickets along themain drainage of the basin.
The high species richness of wet meadows andmoist rock crevices, with 105 species present,makes for a large number of indicator species.The most important of these for wet meadows areSenecio triangularis, Calamagrostis canadensis,Carex spectabilis, C. nigricans, Vaccinium nivic-tum, Aster alpigenus, Dodecatheon jeffreyi, D.subalpinum, Eriophorum criniger, Juncus merten-
2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 187
sianus, and the moss Polytrichum juniperinum.Areas with saturated soils in early summer oftensupport stands of tall herbs such as Veratrumcalifornicum, Mertensia ciliata, Sphenosciadiumcapitellatum, and Helenium bigelovii.
Although there is a significant overlap of wetmeadow species with those of dry meadows, acondition not surprising in view of communitygradients of moisture availability and soil depth,the high cover of wet meadow associations isoften but not invariably a distinguishing feature.In some areas, wet meadows grade gradually intowillow thickets, while other areas exhibit atransitional association between wet and drymeadows formed by a heather turf occurring inthe granitic joint along the east face of the basin.This latter community is dominated by Phyllo-doce breweri, with Vaccinium nivictum, Juncusparryi and several dry meadow herbs as associ-ated species. While P. breweri is widespreadacross many habitats in the basin, only in thisarea does it form the dominant groundcover.
Moist rock crevice communities are concen-trated on the east face of the basin, on benchesalong the major joint area, and in various pocketsof rocky talus. These are areas below wetmeadows that provide sufficient moisture inputsto support the growth of mesic shrubs and herbs.While plants cover the total surface along thesemoist crevices, the crevices themselves cover lessthan 5% of the rock surface where they occur.Characteristic species in these habitats includeSambucus racemosa, Lonicera involucrata, Mer-tensia ciliata, Aquilegia pubescens, and Thalictrumfendleri. Although plant cover over the crevicesthemselves is virtually 100%, only about 5% ofthe rock faces provide crevice habitats.
A related habitat includes a group of speciesthat grow among large boulders 1–3 m indiameter in slide areas which maintain moistsoils throughout the summer. In addition to thespecies above Lonicera conjugialis, Aquilegiaformosa and Helenium bigelovii are commonlypresent, while smaller herbaceous perennials arelargely absent.
Dry meadows and dry rock crevices. Drymeadow associations occupy a relatively largearea of the Emerald Lake Basin, most notably inthe relatively flat plateau and benches below thenortheast ridge and boundary of the basin. Inthese sites, soils are formed of shallow layers ofdecomposed granite with low organic matter, andtypically dry early in the growing season asindicated by the early senescence of herbaceousspecies. There are also significant areas of drymeadow northeast of Parson’s Pond, adjacent tothe stand of Pinus monticola in the northeastcorner of the basin, in the northeast fault area,and in small, scattered areas of relatively flattopography on south-facing slopes.
The overall diversity of dry meadow and drycrevice associations is surprisingly high, with 84species encountered. Indicator species for the drymeadow habitats include Ivesia santalinoides,Juncus parryi, Dicentra nevadensis, Eriogonumincanum, Calyptridium umbellatum, Sedum obtu-satum, and grass species such as Achnantherumoccidentale and Muhlenbergia filiformis. Thedensity and cover of dry meadow associations ishighly variable, with moisture relations a criticalfactor (Klikoff 1965; Burke 1982; Benedict 1983).Much of the area of dry meadows has only 20–25% plant cover, although the dry meadownortheast of Parsons Pond has 50% or morecover. Small stands of Dicentra nevadensis highon the east slope of the basin approach 100%groundcover.
Dry rock crevices are widespread all across thefractured areas of granite faces in the EmeraldLake Basin, but highly diffuse in coverage andaccount for only a very small total area. Thesehabitats, usually relatively narrow cracks nomore than 50 cm wide, support a number ofspecialist species as well as herbaceous perennialscommon to dry meadows. While these crevicesappear dry at the surface, the presence ofsuffrutescent perennials suggests good wateravailability deep in the crevices. The characteris-tic dry crevice species include Penstemon new-berryi, Holodiscus microphyllus, Spiraea densi-flora, and Sedum obtusatum. Eriogonum nudum,Ivesia pygmaea and a variety of dry meadowherbs may also be present but are eclipsed incoverage by the larger shrubs. As with the wetrock crevices, plant cover is often close to 100%along these cracks, but these crevices cover nomore than 5% of the rock surface.
Colluvium. The colluvium association occurs ina few steeply sloped areas along the fault trace onthe southwestern boundary of the basin and intwo smaller areas to the south where largeboulders change to smaller granitic boulders0.3–1.0 m along their major axis. The 26 speciespresent occurred in areas with shallow soildevelopment between the boulders. Most of thehighly heterogeneous plant cover, which averagedabout 25% of ground surface, was composed ofsmall herbaceous perennials up to 20 cm inheight. Species which dominated this habitat bothin abundance and cover were Senecio fremontii,Eriogonum incanum, Carex lanuginosa, Phloxdiffusa, and Erysimum capitatum. The first ofthese is particularly important, making up amajor part of the cover. Two pteridophytes,Cryptogramma acrostichoides and Selaginellawatsonii, grow along the margins of the bouldersand provide significant cover. Two species,Primula suffrutescens and Anaphalis margarita-cea, appear to be restricted to the colluviumhabitat.
188 MADRONO [Vol. 56
Fellfields. Relatively little area of alpine fell-field is present in the Emerald Lake Basin. Thiscommunity is found at high elevation on aplateau below Alta Peak, a site with Pinusbalfouriana on a north-facing slope overlookingPear Lake. Compared to the colluvium habitat,the fellfield has a ground surface scattered withsmaller granite rocks and a large area of exposedsoil. Unlike the colluvium habitat where plantshave colonized virtually all of the available areasof bare soil, this fellfield is very sparsely vegetatedand low in diversity. Nowhere is plant covergreater than about 10%, with a typical covercloser to 5%. Seven species were encounteredhere—Chaenactis alpigena, Carex helleri, Ranun-culus escscholtzii, Minuartia nuttallii, Eriogonumincanum, Saxifraga tolmiei and Trisetum spica-tum. The first two of these were encounterednowhere else in the basin.
DISCUSSION
The relatively large species diversity of theEmerald Lake Basin is consistent with similarlevels of diversity reported for other high SierraNevada basins. Burke (1982) reported the pres-ence of 277 plant species in the Rae Lakes Basinwhich covers approximately 6 km2 with eleva-tional ranges from 3046–4040 m. Cheng (2004)provided summary descriptions of U.S. ForestService Research Natural Areas in the centralSierra Nevada that included subalpine meadowand alpine talus and scree slope communities.Three of these, Clark Fork (1869–3826 m eleva-tion, 874 ha), Highland (2650–2815 m elevation,178 ha), and Snow Canyon (2499–3001 m eleva-tion, 285 ha) contained 227, 200, and 223 taxa,respectively. These levels of biodiversity can beexplained only partially by the elevational gradi-ents included within the basins. More importantis the diversity of habitat conditions presentdespite a low level of plant cover and biomass.
The relative abundance of woody species in theEmerald Lake Basin provides evidence of the lessstressful conditions of this subalpine area com-pared with true alpine communities where shrubspecies are rare (Rundel et al. 2008). Neverthe-less, herbaceous perennials do form the largestlife form in the flora as they do in alpine habitats.In both subalpine and alpine habitats, herba-ceous perennials have the characteristic ofmaintaining large proportions of total biomassbelowground where they play an important rolein carbohydrate storage over the winter months(Mooney and Billings 1960; Billings 1974; Rundelet al. 2005).
Interesting among the Poaceae in the EmeraldLake Basin is the presence of three species ofMuhlenbergia, grasses with C4 metabolism. Theseinclude two perennial and one annual species.Although C4 metabolism in grasses is normally
characteristic of low, subtropical habitats, Muh-lenbergia forms an exception with species reach-ing to alpine environments (Sage and Sage 2002).
The 7% of the flora represented by annualplant species at Emerald Lake is similar to typicallevels of about 8% reported for the summer dryalpine areas of the Sierra Nevada and WhiteMountains (Jackson 1985; Jackson and Bliss1982; Rundel et al. 2008). Annual species are rarein typical circumboreal arctic-alpine floras of theNorthern Hemisphere where they commonlycomprise only 1–2% of the flora (Billings 2000).
Ecological and biogeographic data on high-mountain and related alpine meadows in theSierra Nevada have recently been summarized(Fites-Kaufman et al. 2007; Sawyer and Keeler-Wolf 2007), as has the general state of knowledgeon Sierra Nevadan meadows in an earlier report(Ratliffe 1985). The diverse associations ofsubalpine meadow communities in the EmeraldLake Basin illustrate the complexity of adaptingsimple systems of community classification asproposed in previous studies for specific basins(Pemble 1970; Burke 1982; Benedict 1983; Taylor1984).
Clearly soil depth and the amount andseasonality of soil moisture availability arecritical components of species distribution. Adetailed study of the floristics and physiographicclassification of seven subalpine meadows inSequoia National Park found that these factorsas well as soil frost action and conditions of waterchemistry, temperature, and depth of floodingexplained the major part of floristic variation(Benedict and Major 1982; Benedict 1983).Subalpine dry meadow associations have beendescribed for other areas of the Sierra Nevada(Klikoff 1965; Ratliff 1979, 1982; Taylor 1976;Burke 1982), but with only limited understandingof controlling factors for species distribution.
The plant communities of the Emerald LakeBasin would fall within a variety of associationsas described in the classification system of Sawyerand Keeler-Wolf (1995). Lower areas of the basinwould be classified as Mixed Subalpine Forest,while associated ericaceous shrublands and wil-low thickets would be termed the MontaneHeather–Bilberry, Montane Wetland Shrub, orHolodiscus Series. The majority of the basinwould fall under ‘‘habitat’’ classifications of thissystem as Subalpine Upland Shrub habitat,Subalpine Wetland Shrub habitat, SubalpineMeadow habitat and Alpine habitat. Thesecategories are much too crude, however, toappropriately separate the plant associationspresent in the Emerald Lake Basin.
With continuing concerns about global change,nutrient enrichment, and acid deposition in highmountain basins such as that of Emerald Lake, itis important to have baseline data on floristicdiversity and ecological communities. Continuing
2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 189
studies in the Emerald Lake Basin can provide ameans of identifying and assessing future envi-ronmental changes.
ACKNOWLEDGMENTS
We thank the California Air Resources Board forsupporting this research. Gail Baker, Evan Edinger,Carmen Crivellone, and Karen Poulin provided majorassistance in carrying out the field work for this project.The assistance of Kathy Tonnessen, Dave Parsons, andthe staff of Sequoia and Kings Canyon National Parksis acknowledged with thanks.
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2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 191
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CO
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.2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 193
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X
AP
PE
ND
IX1.
CO
NT
INU
ED
.
2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 195
Div
isio
n/F
am
ily
Sp
ecie
so
rin
frasp
ecif
icta
xo
nL
ife
form
Dry
fore
stW
illo
wth
ick
etW
etm
ead
ow
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crev
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mea
do
wD
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evic
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ela
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ier
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A.
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A.
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nti
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mm
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m.
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sp.
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on
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HP
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orm
skjo
ldii
Ro
emer
&S
chu
ltes
HP
X
AP
PE
ND
IX1.
CO
NT
INU
ED
.196 MADRONO [Vol. 56
Div
isio
n/F
am
ily
Sp
ecie
so
rin
frasp
ecif
icta
xo
nL
ife
form
Dry
fore
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wth
ick
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etm
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crev
ice
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mea
do
wD
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yd
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ene
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son
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tle
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ary
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ven
enosu
sS
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ats
on
HP
-GX
AP
PE
ND
IX1.
CO
NT
INU
ED
.
2009] RUNDEL ET AL.: EMERALD LAKE BASIN VEGETATION AND FLORA 197
Div
isio
n/F
am
ily
Sp
ecie
so
rin
frasp
ecif
icta
xo
nL
ife
form
Dry
fore
stW
illo
wth
ick
etW
etm
ead
ow
Wet
crev
ice
Dry
mea
do
wD
rycr
evic
eC
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uviu
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ellf
ield
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ace
ae
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nath
eru
mle
mm
onii
(Vase
y)
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wo
rth
HP
XA
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identa
le(T
hu
rber
)B
ark
wo
rth
HP
XX
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stis
idahoensi
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ash
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urb
eri
ana
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chc.
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abil
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yd
b.
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mus
cari
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k.
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HP
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um
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ner
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der
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ark
wo
rth
HP
XT
rise
tum
spic
atu
m(L
.)R
ich
ter
HP
XX
XX
AP
PE
ND
IX1.
CO
NT
INU
ED
.
198 MADRONO [Vol. 56
