Relationships Between Scotch Broom (Cytisus Scoparius), Soil Nutrients, and Plant Diversity in the Garry Oak Savannah Ecosystem
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Plant Ecol DOI 10.1007/s11258-009-9655-7 Relationships between Scotch broom (Cytisus scoparius), soil nutrients, and plant diversity in the Garry oak savannah ecosystem Jacqueline Shaben Æ Judith H. Myers Received: 23 May 2007 / Accepted: 14 April 2009 Ó Springer Science+Business Media B.V. 2009 Abstract Scotch broom (Cytisus scoparius), is a broom-invaded plots. Finally, in greenhouse assays leguminous shrub, native to the Mediterranean, which grass growth was not affected as a result of being has invaded most of the remaining Garry oak savan- grown with broom; however, grasses appeared to nah ecosystems in Oregon, Washington, and British produce more flowers when grown with broom. We Columbia. Here, it is considered to be a threat to the conclude that broom does not necessarily modify soil native plant community. We tested the hypothesis that nitrogen availability but may deplete soil phosphorus broom would increase available soil nitrogen by availability and that broom invasion can be associated comparing soil nutrients in contiguous broom-invaded with increase of exotic species and/or the decline of and non-invaded sites. We then looked for changes in native species. patterns of diversity in the herbaceous community that might indicate a role of Scotch broom in changing Keywords Scotch broom Á Cytisus scoparius Á conditions following its invasion. Finally we carried Garry oak savannah ecosystem Á PRSTM probes Á out greenhouse assays to test whether broom had a Soil nitrogen Á Invasive plant greater impact on the growth of a native and an introduced grass compared to that of a native shrub. Broom was associated with only a weak trend in Introduction increased soil nitrogen, but a significant decrease in soil phosphorus was observed. Patterns of plant Scotch broom (Cytisus scoparius (L.) Link), a legu- diversity differed between two sites. At one site, minous shrub native to the Mediterranean, has 60% of the plants whose abundances increased in the become an invasive plant of ecological concern in broom-invaded plots were introduced species while several areas of the world, including parts of Australia native species abundances decreased in the broom- (Downey and Smith 2000), New Zealand (Williams invaded plots compared to broom-free plots. At the 1981), and North America (Bossard and Rejmanek other site, 60% of the plants that caused the differ- 1994; Prasad 2003). Broom is well adapted to a ences between broom-invaded and un-invaded plots moderately dry, temperate climate and is intolerant of were native species that were less abundant in the shading by trees and other shrubs. Its invasive nature is attributed to prolific seed production, persistent seeds, year-round photosynthesis due to photosyn- & J. Shaben ( ) Á J. H. Myers thetic stems, production of alkaloid compounds, Department of Zoology, University of British Columbia, 6270 University Blvd, Vancouver, BC V6T 1Z4, Canada formation of dense stands, a lack of natural enemies e-mail: [email protected] in new environments, and the ability to fix nitrogen 123 Plant Ecol (Williams 1981; Wink et al. 1983; Wheeler et al. speculate that the establishment of D. glomerata is 1987; Waterhouse 1988; Prasad 2003; Downey and facilitated by increased nitrogen availability as a Smith 2000). While broom frequently invades dis- consequence of broom invasion, and that this results turbed sites in British Columbia (BC), Washington, in a decline in native plant species and a change in and Oregon, its successful invasion of intact frag- community structure. Fertilization experiments in old- ments of the Garry oak savannah ecosystem (GOSE) fields with D. glomerata by Gurevitch and Unnasch (Parker 2001), an endangered oak and grass habitat (1989) demonstrated such a change in community (COSEWIC 2000; Fuchs 2001) makes it a species of structure. particular conservation concern. The objective of this study is to determine some of Like many legumes, broom has symbiotic root- the impacts of broom on the ecology of native plants associations with nitrogen-fixing bacteria (Rhizobia of the GOSE in southern British Columbia and to sp.). For example, Fogarty and Facelli (1999) deter- contribute to the management of the small fragments mined that broom-invaded soils in forests of South of this ecosystem that remain in order to optimize - Australia had higher levels of nitrate (NO3 ) and native plant species conservation. Here, we evaluate ? ammonium (NH4 ) than did adjacent non-broom five aspects of the impacts of broom in the GOSE by soils, and they suggested that broom is probably (1) Quantifying the seasonal soil nutrient availabil- responsible for changing soil nutrient availability. The ity correlated with Scotch broom-invasion in the ability to fix atmospheric nitrogen allows broom GOSE of southern Vancouver Island. to establish in sites with nutrient-poor soils, likely (2) Comparing the level of soil nitrogen and phos- affording it a competitive growth advantage over non- phorus availability in locations with and without N-fixing plants, and thus this is cause for concern in Scotch broom. the management of GOSE. The negative effects of (3) Determining if the level of soil nitrogen avail- nitrogen-fixing shrubs on invaded plant communities ability and plant community structure are related has already been well established in other ecosystems, in these GOSE sites. such as young volcanic sites in Hawaii (Vitousek and (4) Determining if plant diversity differs between Walker 1989; Hughes and Denslow 2005), Califor- broom-invaded and un-invaded plots in the nian coastal dune ecosystems (Maron and Connors GOSE, and whether plant community differ- 1996; Pickart et al. 1998), grassland plains of southern ences are created by changes in native and exotic US and northern Mexico (Hibbard et al. 2001), and in plant abundances. the South African fynbos (Stock et al. 1995). (5) Ascertaining whether or not an exotic nitrophyl- One way in which increased soil fertility can alter lic forage grass benefits greater than a native plant community structure is by facilitating the inva- grass when grown in association with broom in sion of exotic, competitive species such as nitrophyllic greenhouse experiments. forage grasses (Vitousek and Walker 1989;Weiss 1999). These grasses are particularly good at acquiring available soil nitrogen which allows them to grow quickly and compete successfully with other plants Methods for water, light, and other nutrients (D’Antonio and Vitousek 1992). The dense, copious leaf litter produced Sites by these grasses can alter the microclimate by atten- uating sunlight, insulating the soil from temperature To determine whether Scotch broom invasion is fluctuations, and moderating soil moisture resulting in correlated with higher levels of plant-available nitro- altered decomposition rates and possibly decreased gen, we sought study sites at which broom was still seed germination of native plants (Facelli and Pickett invading a GOSE location. This proved difficult as 1991). most sites were either completely invaded by broom, In the Garry oak savannah ecosystem, Dacty- had previously been invaded, and were now managed lis glomerata (L.) is one of the most common exotic, for no broom or had no broom at all. Two locations invasive forage grasses. GOSE site managers with contiguous areas invaded and un-invaded by 123 Plant Ecol broom and with similar slope, aspect, and substra- Soil nutrient measurements tum were eventually identified in the GOSE on southern Vancouver Island: Rocky Point (48°19.5000 Soil nutrient measurements were conducted using two N, 123°32.5500 W) and Bamberton (48°34.7600 N, methods: standard extraction from soil core samples 123°31.3000 W) (Fig. 1). to obtain the background total nutrient content, and At Rocky Point, five plots were established within a ion-exchange membrane probes to obtain in situ Garry oak savannah that had been used historically by available nutrient levels. the First Nations people for harvesting camas bulbs In February 2005, 15 soil cores were taken (20 mm (Camassia quamash (Pursh) Greene) and were peri- diameter, 0–10 cm depth, including organic and odically burned. More recently, European immigrants mineral soil) from each broom and non-broom plot used the site for sheep grazing (Arthur Robinson, at both sites. These were pooled, air-dried for 72 h and personal communication, 2006). All five plots at sieved through a 2 mm sieve. Soils were then analyzed Rocky Point had negligible slope, were south-facing, for pH, % C, total Kjeldahl nitrogen (TKN), avail- and were between 50 and 80 m apart. able P, Ca and K. We calculated the carbon to nitrogen The Bamberton property was only large enough to ratio (C:N) and % organic matter (from total C 9 support three study plots. This undisturbed site 1.724). received very little human traffic. It was situated From January 22, 2005 through June 12, 2005, directly south of an abandoned cement plant, which ion-exchange membrane (Plant Root Simulator, we later discovered had an impact on soil nutrients PRSTM) probes were used to determine soil nutrient reflected in high levels of calcium. The three plots availability during the course of the growing season. were east-facing and on a steep (60%) slope. Each pair of probes consists of one anion-exchange Fig. 1 Map showing locations of the two study sites on Southern Vancouver Island. Bamberton is south of Duncan, BC, on the east side of the Malahat Highway. Rocky Point is on Department of National Defense property in Metchosin, BC 123 Plant Ecol membrane probe and one cation-exchange membrane Separate t-tests were used to compare richness of probe. Probes work by adsorbing ions as they become plant species (species number) and evenness of plant available in the soil; therefore, the unit of measure- abundances (Pielou’s J’) by treatment. ment is a rate of ion availability and is described in To ascertain which species were responsible for mass of nutrient per membrane surface area per time driving the differences seen between the broom and buried.