Eastern Washington University EWU Digital Commons
EWU Masters Thesis Collection Student Research and Creative Works
2013 Ecohydrology effects of an invasive grass (Phalaris arundinacea) on semi-arid riparian zones Adam D. Gebauer Eastern Washington University
Follow this and additional works at: http://dc.ewu.edu/theses
Recommended Citation Gebauer, Adam D., "Ecohydrology effects of an invasive grass (Phalaris arundinacea) on semi-arid riparian zones" (2013). EWU Masters Thesis Collection. 236. http://dc.ewu.edu/theses/236
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"###! $"%&!'(&!)"*+,-.%/!0/-*1./0/23*!4056*7!-3!-88!*"3/*!-29!*-0:8"2%!9-3/*!3,.;1%,;13! 3,/!<='"2%!*/-*;2&!???????????????????????????????????????????????????????????????!@(! ! $"%!'A&!B,-2%/!"2!%.;129>-3/.!8/C/8*!-3!3>;!B;>!B.//D!*"3/*&!???????????????????????!@A! ! $"%!'E&!B,-2%/!"2!%.;129>-3/.!8/C/8!-3!3,.//!B.-F!B.//D!*"3/*&!??????????????????????!@E! ! $"%!'G&!B,-2%/!"2!%.;129>-3/.!8/C/8!-3!3>;!H-3-,!B.//D!3."F13-."/*&!???????????????!@G! ! $"%!<=&!B,-2%/!"2!%.;129>-3/.!8/C/8!-3!3>;!I;+D!B.//D!*"3/*&!???????????????????????!(=! ! ! "#! Introduction The transitional zones between terrestrial and aquatic habitats provide many ecological functions such as flood control, water filtration, and high ecological diversity. These riparia are particularly crucial in semiarid regions with limited water resources. Invasive alien species can cause riparian ecosystem function to decline (Junk et al. 1989, Gregory et al. 1991, DeFerrari & Naiman 1994). In an effort to quantify this decline my study investigates the ecohydrologic (relation between plants, water, and nutrients in an ecosystem) impacts of the aggressive invader, reed canary grass (Phalaris arundinacea hence forth RCG) in low order streams in the temperate semiarid region of eastern Washington. Riparia provide a link between upland and aquatic ecosystems, mediating energy transfer and nutrient cycling. The proximity to water and the diverse vegetation in riparian zones offers crucial habitat for both terrestrial and aquatic wildlife (Naiman & Decamps 1993, Naiman 1997, and Ward 1998). The presence of dense riparian vegetation acts as a buffer reducing surface flows and filtering pollutants before they enter stream systems (Lowrance et al. 1985, Daniels & Gilliam 1996, Naiman et al. 2005). The vegetation structure strongly influences the hydrology of riparian zones and adjacent aquatic systems (Roden & Ehleringer 2000, Dahm et al. 2002, Cleverly et al. 2006, Chamier et al. 2012). In some cases, such as on the middle Rio Grande, riparian vegetation can account for 20-50% of water lose from the river (Dahm et al. 2002). Invasive species, particularly those such as RCG that create monotypic stands, can have a significant impact on ecohydrology. For example, in four South African watersheds of historically shrub/grassland habitat, Le Maitre et al. (2002) documented about a 6.7% reduction in stream flow due to the invasion of woody: Pinus, Acacia, and Hakea species. Dye & Jarmain (2004) estimated that the removal of one of these invasive species, !"#"$#%&'#()*$$, could reduce evaporation by up to 600 mm of water a year. In the American southwest, the introduction of the woody invasive tamarisk (Tamarix spp.) as a soil stabilizer has increased soil salinity, modified habitat for native plants and animals and had a significant effect on hydrology (Brotherson & Field 1987). Tamarisk can alter flood cycles by trapping sediment, changing stream slope, and reducing stream channel width, thereby changing the geomorphology of the riparian zone and reducing access to groundwater for native vegetation (Brotherson & Field 1987, Zavaleta et al. 2001, Pataki et al. 2005, Cleverly et al. 2006). Transpiration rates at the leaf scale and evapotranspiration rates at the plot scale are used to measure the amount of water lost from vegetation to the atmosphere, which can indicate the ecohydologic effect of plants. Evapotranspiration of riparian plants, particularly invasive plants, can have significant effects on shallow water tables and stream flow in arid regions (Cleverly et al. 2006, Pataki et al. 2005). Woody vegetation has been the focus for most research investigating the relationship of invasive riparian species’ evapotranspiration rates and their effects on groundwater and stream flow (e.g. Brotherson & Field 1987, Cleverly et al. 2006, Le Maitre et al. 2002). For example, Cleverly et al. (2006) found that as water tables lowered, Tamarix chinensis increased evapotranspiration rates by up to 50% while native species evapotranspiration remained constant. Evapotranspiration rates and water sources can vary widely between plant species (Scott et al. 2000, Dahm et al. 2002). For example, Scott et al. (2000) used ! "! evapotranspiration rates and water fluxes to estimate the water source of two non- phreatophyte species growing in semi-arid riparian zones of Arizona. A grass (Sporobolus wrightii) relied on recent precipitation in shallow soils, while a shrub (Prosopis velutina) used deeper soil waters. In a multi-scale meta-analysis of evapotranspiration rates of native and invasive species, Cavaleri and Sack (2010) found that at the leaf scale, invasive species had higher rates of stomatal conductance and water loss than native plants. Study results varied across different vegetation types at the ecosystem scale, but evapotranspiration rates tended to be higher in systems with invasive species in semi-arid grasslands and systems dominated by herbaceous invaders versus herbaceous natives (Cavaleri and Sack 2010). The small number of studies reviewed (n < 5) for this meta-analysis indicated the need for more research. While transpiration and evapotranspiration rates can indicate the amount of water used, the analyses of naturally occurring stable isotope ratios of hydrogen and oxygen in source water and plant xylem water can add information about the ecohydrology of a system by identifying a plant’s use of water sources (Dawson & Ehleringer 1991, Busch et al. 1992, Ehleringer & Dawson 1992, Dawson et al. 2002). The ratio of heavier isotopes (18O and 2H, which is often abbreviated D for deuterium) to lighter isotopes (16O and 1H) in a water sample as compared to that of the Standard Mean Ocean Water (SMOW; ratio denoted as ! values) may allow for water sources to exhibit distinct isotopic ratios. !D= ((D/H) sample /(D/H) SMOW -1) x 1,000‰ These differences in ! values of water sources indicate fractionation which is due to thelower mobility of heavier isotopes relative to the lighter isotopes. The amount of ! "! fractionation changes with temperature associated with changes in altitude, latitude, and seasonal recharge/depletion. Generally speaking, the change in seasons from warmer to cooler temperatures reflects a depletion of the heavier O and H isotopes in water, resulting in a more negative ! value. Similarly, lower temperatures with increases in altitude and latitude are associated with a lower ! value (Dawson et al. 2002). Within a general location, surface water sources can experience more recent inputs and greater evaporation than deeper water while deeper water sources can contain a mixture of past seasonal inputs resulting in between shallow and deep-water sources different isotopic ratios (Dawson et al. 2002). Plant tissue water that has not been exposed to evaporative processes during water uptake by the roots, such as those found in the xylem and branches, will have an isotopic signature indicating the relative proportions of its water sources (Ehleringer & Dawson 1992, Dawson et al. 2002). Cordell and Sandquist (2008) used isotopic ratios to determine the effects of an invasive grass on the water use of a native tree species in a Hawaiian dry land forest. They found that in the absence of the invasive grass, native trees used higher proportions of shallow water sources and exhibited morphological characteristics associated with reduced water stress. Using stable isotope ratios of oxygen, Busch et al. (1992) found that tamarisk switched water sources from shallow to deeper sources in times of low water while native tree species use only deep water source, giving tamarisk a competitive advantage in drought conditions. While there is substantial research documenting invasive woody species’ effects on hydrology in riparian zones, there is increasing evidence that grass can also alter the hydrology of a system. Both Schilling & Kiniry (2007) and Wilcox et al. (2008) found ! "! that grasses can have evapotranspiration rates comparable to woody vegetation, whereas a meta-analysis by Cavaleri and Sack (2010) found invasive grasses to have higher rates of primary productivity and evapotranspiration than native species. Watts and Moore (2012) found the invasive riparian grass, Arundo donax to have plot scale transpiration rates of 8.8 mm water per day, exceeding rates found by Cleverly et al. (2002) for tamarisk of 7 mm a day. Whether woody or herbaceous, the increase in water use by invasive species compared to natives alters the echoydrology of riparian systems, potentially causing streams to experience below historic low flow conditions. Reduced flow allows higher concentrations of pollutants to develop (Lee 1969, Allan and Castillo 2007, Carroll 2007, Tarbutton et al. 2010) and contributes to poor habitat for salmonids and other aquatic life through higher water temperatures and reduced dissolved oxygen. Reducing the amount of water in an aquatic system reduces habitat area by decreasing flow rates needed for spawning and prey location, and decreasing open water areas for migrating waterfowl (Mantua et al. 2010). Many streams in eastern Washington have been experiencing below average low flow rates. For example, Crab Creek has experienced a reduction in monthly average discharge (Fig. 1). Many water bodies in the semiarid region of the Palouse Prairie and Channeled Scablands of eastern Washington do not comply with standards set by the federal Clean Water Act. These streams have elevated levels of fecal coliform bacteria, decreased dissolved oxygen, altered pH, and increased water temperature (e.g. Carroll 2007, Joy et al. 2009, Tarbutton et al. 2010) exacerbated by low summer stream flows. There are likely many causes of below historic average flows in semiarid regions: climatic effects, increased water use through urbanization and agriculture, and, as I ! "! investigated, changes in vegetation structure. RCG is one species that could have a significant impact on the hydrology of stream systems. RCG is a perennial, long-lived, C3, cool season grass species that commonly invades riparian zones in the temperate United States, frequently forming dense monotypic stands (Lavergne & Molofsky 2004). It grows 1-2 m tall with dense crowns and vigorous rhizomes that exhibit aggressive vegetative spread (Hitchcock 1950). RCG has genotypes native to the Midwest of North America and Eurasia (Lavergne & Molofsky 2004), but the invasive form appears to be a cultivar of largely Eurasian origin (Lavergne & Molofsky 2007). RCG has had multiple introductions from Europe since 1850 (Merigliano & Lesica 1998). The first introductions were low alkaloid forage crop cultivars. Cultivars have also been actively planted in wetlands and riparian zones to restore degraded soils and waters, including wastewater treatment, by extracting soil contaminants and stabilizing shoreline (Anderson 1961, Apfelbaum and Sams 1987, Lavergne & Molofsky 2004). Recently, RCG has been used as a bioenergy crop and for pulp, paper, and fibers (Casler et al. 2009, Lavergne & Molofsky 2004). Due to these multiple introductions RCG is currently established in most of temperate North America (Hitchock 1950, Galatowitsch et al. 1999). This has led to an increase in both the genetic diversity and the phenotypic plasticity of the species with invasive genotypes of this species emerging earlier in the season and showing increased tillering rates, leaf production, and total biomass production (Lavergne & Molofsky 2006). These factors have promoted the invasion of Eurasian strains of RCG across a variety of ecosystems and resource gradients. ! "! RCG can form dense monotypic stands in wet prairies, riparian areas, and some uplands, reducing biodiversity (Barnes 1999, and Paveglio & Kilbride 2000). When occupying riparian areas, RCG can increase sediment deposition and reduce water circulation through the production of dense vegetative stands that grow into the stream channel (Hudgson 1968, Werner & Zedler 2002). Lavergne & Molofsky’s 2004 review cited several studies that found RCG to dominate 50-100% of the areas where it was introduced. For example, Tanner et al.’s 2002 study in Everett, Washington, found this grass to occupy 66% of the wetlands on a freshwater intertidal island and Barnes (1999) found RCG to occupy 40% of a Wisconsin stream within 15 years of introduction, indicating that this species is a rapid and dominant invader. The plasticity of RCG enables it to be productive under a wide range of moisture levels including drought and flood conditions (Figiel et al. 1995, Miller & Zedler 2003, Morrison and Molofsky 1998). Miller & Zedler (2003) compared two plants that occupy similar habitat, Spartina foliosa (California cordgrass) and RCG, under different hydroperiods. They found that flooding decreased allocation to below ground biomass but increased shoot length in RCG. When grown alone these species produced similar above ground growth but when grown together RCG’s above ground growth was twice that of California cordgrass. This allowed RCG to occupy a greater cover area than California cordgrass in their study plots, increasing RCG’s light capturing ability. Under increased flooding, RCG formed tussocks, allowing it to more than double its above ground biomass (Herr-Tutoff & Zedler 2007). Its adaptations to different water regimes allow RCG to out compete other herbaceous riparian and wetland plants. ! "! RCG can withstand high nutrient and sediment loads. In a study with eight wetland species, RCG had higher establishment rates as nitrogen and sedimentation increased (Mahaney et al. 2004). Kercher & Zedler (2003) found RCG to increase productivity by 35% and 195% under low and high nutrient levels respectively. They also found multiple disturbance factors work synergistically to increase invasion of RCG. In mesocosm studies comparing RCG with native species, RCG increased root/shoot ratio with decreased nutrients. RCG has been found to increase its overall biomass, reduce root allocation (Green and Galatowitsch 2001) and increase clonal spread and tiller production (Mauer & Zedler 2002) with increased nutrients (Wetzel & van der Valk 1998). Many studies have looked at possible methods for control of RCG including biocontrol, grazing, herbicide, and disking (Lavergne & Molofsky 2006). In testing a biocontrol insect, Hansen et al. (1985) found that, of 16 grass species, RCG was among the least favored by black grass bug (Labops hesperius). RCG biomass has been show to decrease with rotational and continuous grazing and to increase with exclosures (Paine & Ribic 2002). Various chemical controls have been used for RCG with limited long-term success without additional and repeated control methods (Apfelbaum & Sams 1987, Annen et al. 2005, and Lavergne & Molofsky 2006). In southwestern Washington, applications of herbicide have been combined with mechanical disking over three growing seasons, resulting in reduced stem density of RCG in wetlands (Kilbride & Paveglio 1999, Paveglio & Kilbride 2000). Some of the most effective control methods have used multiple control methods combined with establishing overstory tree communities, producing shade to reduce RCG biomass (Kim et al 2006, Miller et al. 2008) ! "! Although RCG has been well studied, little is known about its water use or how that affects the ecohydrology of riparian zones. Schilling and Kiniry (2007), using a combination of field observations and model simulations calculated that the average water use of RCG was 3.3 mm/d in July and 2.3-2.8 mm/d May, June, August, and September. Their study used field measurements of groundwater fluctuation to create a model of vegetative water use. Unlike many woody riparian species, there has been little in situ research on the direct water use of RCG, how that compares with other riparian species, and its hydrologic effects on an ecosystem. The goal of my study is to determine RCG’s effects on the ecohydrology of semi- arid riparian zones. I hypothesis that compared to other riparian vegetation, RCG will have a longer growing season, have greater cover in the riparian zone, have both higher water use at the leaf scale and plot scale than other species, and use shallow water sources with a close hydraulic link to the stream. These factors will allow RCG to have a significant impact on the ecohydrology in these semi-arid riparian zones. ! "! !"#$%&'( ! )#*&+(,-".( ! My study sites are located in the Palouse Prairie and Channeled Scablands of semiarid eastern Washington (24.5-50.8cm of annual precipitation). The northern extent of the Palouse Prairie bioregion overlaps with the Channeled Scablands where areas of higher elevation retained the highly productive loess soils characteristic of the Palouse Prairie. The Channeled Scablands were formed when successive Ice Age glacial floods scoured 24,000 square kilometers of basalt bedrock overlain with a deep layer of windblown loess (Bretz 1928, Bretz 1969). These massive floods eroded canyons into the basalt bedrock, forming many drainage channels, coulees, and potholes. The combination of these two landscape features created an environment suitable for wheat production, grazing, and urbanization within several watersheds. Riparian zones in this region include a mix of climax shrub communities consisting of snowberry (Symphocarpos alba), black hawthorn (Crataegus douglasii), and woods rose (Rosa woodsii) and of forest consisting of ponderosa pine (Pinus ponderosa), cottonwood (Populus trichocarpa), quaking aspen (Populus tremuloides), and red alder (Alnus rubra: Black et al. 2003, Crawford 2003). They also provide breeding, rearing, and refuge habitat for many species of wildlife including elk, moose, white deer, mule tail deer, and the endangered sage grouse (Centrocercus urophasianus: Connelly et al. 2000). My specific $%&'(!$)%*$!+*,*!+)%-).!/0&,!+1%*,$-*'$!23145*!"6!7)8!9:;!Crab, Cow, Rock, and Latah Creeks. Crab Creek is 163 km in length and drains approximately ! "#! 8203 sq km from Reardan, WA to the Columbia River. My three Crab Creek sites are located in Lincoln country on BLM land approximately 26-39 km east of Sprague, WA. Cow Creek has ephemeral flows for 80 km starting at Sprague Lake and draining into the Palouse River. My two Cow Creek sites are just south of Sprague Lake in Adams County. Rock Creek intermittently flows south from Turnbull National Wildlife Refuge (Spokane County) to Rock Lake (Whitman County). Two sites are located within the Rock Creek drainage; one site is just north of Rock Lake on the Pine Creek tributary and the other site is located near the headwaters on the outflow of Pine Lake within Turnbull National Wildlife Refuge. Latah (Hangman) Creek runs 97 km and drains 1083 sq km from Benewah County, Idaho to the Spokane River in Spokane, WA. My Latah Creek sites are located on two tributaries: Mica Creek and a different Rock Creek both near the town of Rockford, WA. ! ! Overall Experimental Design Nine sites were stratified across the four watersheds, with two or three sites per watershed. The criteria for site selection included the presence of relatively natural riparian zones (free from human disturbance) with RCG approximate abundance ranging from 10 to 80%. Each of the nine study sites comprises a 200 m stream reach (hereafter study site). Sites were intentionally selected with varying amounts of RCG to compare effects of different amounts of RCG.! !! ! ! ! ""! !"#"$%$&'()%(*)+"','-./&0)12-3"4! ! $%&''!(&)*+',(+!-'&'!&)*./012!1/,)('.!-3(%3*!'),%!+(4.2!+3('!(/!)++'++!5/(%! 6)+,41)&!71)*(!,/004*3(3'+!)*.!8'/0/&7%3,!7/+3(3/*+9!$&)*+',(+!-'&'!1/,)('.! 7'&7'*.3,41)&!(/!(%'!+(&')0!)*.!+(&'(,%'.!),&/++!(%'!'*(3&'!&37)&3)*!:/*'!;(%'!)&')! 3*<14'*,'.!52!+(&')0!<1/-!)(!+/0'!7/3*(!.4&3*8!(%'!8&/-3*8!+')+/*=9!!$%'!&37)&3)*! :/*'!-)+!.'*/('.!52!5/(%!6'8'()(3/*!)*.!8'/0/&7%3,!(&)3(+!;>477!?!@+('&A)07! 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ugas et al. 1993, LI-COR 1999) (-!-2$!0/-'0L!!@MNO!(9:!H$2!H+''P)!! Q'(05+','9-0!2'+'!-(P'9!(-!-.+''!:/%%'+'9-!6'+/$:0!:5+/98!-.'!7R"7!8+$2/98! 0'(0$9L!'(+&J!;,/:!Q(J?A!,/::&'!;'(+&J!S5&J?A!(9:!&(-'!8+$2/98!0'(0$9!;'(+&J! K'6-',3'+?)!!K(,6&/98!2(0!:$9'!-2/4'!:5+/98!'(4.!6(+-!$%!-.'!8+$2/98!0'(0$9!%$+! '(4.!%$450!06'4/'0!;9>T?!6+'0'9-!2/-./9!-.'!8'$,$+6./4!6$0/-/$90)!K$,'!06'4/'0! 2'+'!6+'0'9-!(-!,$+'!-.(9!$9'!8'$,$+6./4!6$0/-/$9!2./&'!$-.'+0!2'+'!4$90-+(/9':! -$!4'+-(/9!6$0/-/$90)!U&&!B"!,'(05+','9-0!2'+'!-(P'9!3'-2''9!-.'!.$5+0!$%!"RLRR! (9:!"VLRR!$9!9'(+!4&$5:&'00!:(J0)!! ! @$!:'-'+,/9'!-.'!./8.'0-!6$-'9-/(&!06'4/'0!-+(906/+(-/$9!+(-'!6'+!59/-!$%! 8+$59:!(+'(!;"G,7?!%$+!'(4.!%$450!06'4/'0!(-!'(4.!8'$,$+6./4!6$0/-/$9A!1!,5&-/6&/':! -.'!S5&J!-+(906/+(-/$9!+(-'!6'+!&'(%!(+'(!3J!-.'!KFU!-/,'0!-.'!3/$,(00!6'+!,7!$%! 8+$59:!05+%(4')!W+$59:!05+%(4'!-+(906/+(-/$9!-+(906/+(-/$9!+(-'!59/-0!(+'!/9!,!,$&'! 7 D7E<, <0)! ! !"#$%&3+4%5$()&01!2& ! To determine the water sources for my nine focus species as well as ponderosa pine, I used the ratios of stable isotopes (denoted as !) of hydrogen (1H and 2H) and ! "#! oxygen (16O and 18O) to distinguish between deep and shallow water sources (per Ehleringer & Dawson 1992 and Dawson et al. 2002). I used ponderosa pine, with its deep root system, as a proxy for an inaccessible deep groundwater source. Water extracted from plant tissue that has not experienced evaporative processes, including xylem water from stems and branches, has an isotopic signature that has not been altered from its source (Dawson & Ehleringer 1992). Therefore, comparing the !D and !18O in plant-xylem water to local surface and deep water sources can indicate the proportion of shallow groundwater versus deep groundwater used by the plant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stream water = [(! Oplant - ! O gw) / (! O stream - ! 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Drainage # Active Channel Bank Floodplain Upland Slope Site 2 km Species #spp/ % % % #spp/ % % % #spp/ % % % m RCG Herb Woody m RCG Herb Woody m RCG Herb Woody Cow A 275.5 20 0.19 31.5 57.0 0.0 0.72 80.8 403.8 0.0 0.69 2.7 156.7 0.0 Cow B 305.5 54 1.18 22.9 30.5 129.8 0.97 82.0 65.3 43.8 1.24 11.4 260.0 13.6 Crab A 1,405.9 25 2.08 84.5 141.1 0.0 0.93 80.8 53.2 0.0 1.05 14.0 220.0 2.9 Crab B 1,450.0 32 3.11 76.6 710.1 60.7 0.48 11.1 131.2 41.1 Crab C 1,809.2 39 1.48 23.8 228.0 400.0 0.86 67.5 75.9 22.2 0.74 9.9 242.8 182.2 Latah A 60.2 29 0.38 51.8 229.9 117.2 0.40 43.8 134.0 24.8 0.25 8.6 128.6 31.2 Latah B 342.2 19 1.00 100.0 33.3 0.0 0.56 43.4 65.8 10.9 1.12 47.9 82.3 72.3 Rock A 100.0 (TNWR) 46 0.58 59.1 40.1 36.2 1.07 48.3 100.7 229.6 1.05 20.4 157.8 255.3 Rock B 933.4 46 0.52 71.6 78.7 39.2 0.47 27.8 109.1 39.1 1.04 0.0 75.4 24.7 Average 742.4 34.4 1.2 58.0 172.1 87.0 0.7 52.7 112.0 41.2 0.9 14.0 161.6 69.3 ! ! 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B+=&! .+.&! ! ! ! />8>@>A! (+.B! ?.+44! ! ! ! CD23$8%! B+4)! -.+.'! G#$%! :;3%! (+'*! .+)(! ! ! ! CD23$8%! (+'4! .+,)! ! ! ! ! ! ! 52#$0#6! -+%#! -+%#! ! ! 8(9#:1<*+)#*! :; ! "#! $%&'(!#)!$%&'(!*+,-!.%'./'%,(0!(1121!3123%4%,+25!*+,-!6)7)!%50!#89!.25:+0(5.(! +5,(1;%' Floodplain Nettle 5.9 1.49 8.94 2.86 Floodplain RCG 96.3 18.24 133.18 59.45 Floodplain Golden Currant 7.8 1.47 10.87 4.65 Floodplain Woods Rose 14.8 3.78 22.91 6.81 Floodplain False Solomon 3.7 0.88 9.64 5.88 Floodplain Canada Thistle 46.1 12.45 72.14 20.03 Upland Slope Nettle 13.1 2.91 19.07 7.16 Upland Slope RCG 18.8 2.39 23.70 13.89 Upland Slope Golden Currant 3.7 1.12 6.18 1.27 Upland Slope Woods Rose 5.6 1.42 8.60 2.68 Upland Slope Snowberry 6.8 1.42 9.91 3.73 Upland Slope False Solomon 19.0 4.41 28.05 10.00 Upland Slope Canada Thistle 10.6 2.48 15.71 5.54 ! 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