RESEARCH ARTICLE Site-Specific Effects of Exotic Annual Grass Control Integrated with Revegetation L. Noelle Orloff, Jane M. Mangold and Fabian D. Menalled ABSTRACT Invasion by exotic annual grass species such as downy brome and Japanese brome has been implicated in ecosystem degradation in much of the western United States and strategies to restore lands dominated by these species are needed. We evaluated integration of herbicide and revegetation to restore old-field and rangeland sites dominated by annual brome species. In split-plot field studies, we compared three imazapic herbicide rates and eight seeding treatments at an old-field and a rangeland site. We evaluated brome, seeded species, and existing vegetation response to our treatments. Herbicide controlled annual brome at both sites, with > 95% decreases in brome biomass in plots treated with the high rate compared to non-sprayed controls. Control of annual brome persisted for two growing seasons at the old-field site and one season at the rangeland site. Annual brome abundance in non-treated control plots at both sites decreased by 80% between the first and second growing season. No seeded individuals established at the old-field site, but they persisted through two growing seasons at the rangeland site. Seeded species establishment at the rangeland site was impacted by both herbicide rate and seed mix, but no clear patterns emerged. Finally, while we controlled brome at both sites, existing desired vegetation increased only at the old-field site, where perennial grass biomass increased by about 30% in plots sprayed with both rates of herbicide. The differing responses of the existing and seeded plant communities across the two sites highlight the importance of integrating site-specific knowledge into restoration plans. Keywords: Bromus arvensis, Bromus tectorum, Conservation reserve program, downy brome, Japanese brome, land use history xotic annual grass invasion has been implicated in cases, long-term dominance by exotic annual grasses leads Eecological damage and change around the globe to the loss of desired perennial species from plant com- (D’Antonio and Vitousek 1992). In the western United munities through decreased recruitment and seed bank States, downy brome (Bromus tectorum) invasion in par- impoverishment, hindering the long-term success of con- ticular has been associated with decreased recruitment trol programs (Humphrey and Schupp 2001, 2004). These of desired species, altered fire regimes, and degradation differences in plant community responses to management of wildlife habitat (Mack 2011, Balch et al. 2013). This are driven by site-specific factors such as perennial plant species has also been identified as a barrier to restoration community composition, land use history, and abiotic char- efforts in degraded lands (Allen 1995, Di Tomaso 2000). acteristics such as episodic events of episodic precipitation Japanese brome (Bromus arvensis) is another exotic annual for a given year (Radosevich et al. 2007). grass closely related to downy brome (Baskin and Baskin In some cases, weed control and revegetation should 1981). Although Japanese brome does not have the invasive be integrated as a long-term strategy to suppress exotic potential of downy brome, these two species often grow annual grasses and provide habitat for livestock and wild- together in the western United States, particularly in the life (Elseroad and Rudd 2011, Hirsch-Shantz et al. 2014). Northern Great Plains, and are a concern in both cropland However, re-establishing desired species is often difficult on and rangeland settings (Ogle et al. 2003, Gasch et al. 2013). lands dominated by invasive annual grasses (Allen 1995, Di Simply controlling exotic annual grasses does not nec- Tomaso 2000). Though many factors can influence estab- essarily restore a desired perennial plant community on lishment of seeded perennial species, competition from a given site. In some cases, desired perennial vegetation exotic annual grasses in particular can decrease establish- increases in abundance following removal of annual grasses ment of perennial grasses and contribute to failure of seed- (Davies and Sheley 2011, Elseroad and Rudd 2011). In other ing efforts (Hull and Stewart 1948, Romo and Eddleman 1987, Davies et al. 2010, but see James and Svejcar 2010). Ecological Restoration Vol. 33, No. 2, 2015 Suppression of annual grasses may help perennial species ISSN 1522-4740 E-ISSN 1543-4079 to establish during revegetation (Morris et al 2009, Davies ©2015 by the Board of Regents of the University of Wisconsin System. and Sheley 2011). June 2015 ECOLOGICAL RESTORATION 33:2 • 147 Figure 1. Annual and long term precipitation data for the closest weather station to each study site (Ft. Assiniboine, Montana, US for the old-field site [~30 km from site]; Malta 7 E, Montana, US for the rangeland site [~ 8 km from site]). All years in the period of record were used to calculate means (94 and 39 years for Ft. Assiniboine and Malta 7 E, respectively) (WRCC 2014). Imazapic is an herbicide that can be used to control Further, we tested the performance of three species in winter annual grasses in rangelands and old-field sites monoculture and mixes commonly used in revegetation (Morris et al. 2009, Mangold et al. 2013) and help revegeta- settings. We hypothesized that controlling annual brome tion efforts. Integrating imazapic with seeding can favor species would increase seeded species emergence and seeded species, but it could also result in non-target effects. establishment and increase existing perennial plant abun- For example, research suggests that high rates of ima- dance. Second, we hypothesized that the responses of zapic reduce seeded species establishment and damage brome and the existing perennial plant community to our beneficial plants, though these effects are species-specific treatments would be site-specific. and depend on timing of herbicide application relative to seeding (Shinn and Thill 2004, Sheley et al. 2007, Sbatella Methods et al. 2011). Thus, evaluation of perennial species that are competitive with annual grasses and tolerant of imazapic We conducted this study over three years (2009–2011) at rates recommended for annual grass control is needed an old-field and a rangeland site in north-central Mon- to improve revegetation practices. To achieve this goal, tana, U.S.A. Sites differed in land use history, existing we evaluated the integration of herbicide (imazapic) and plant community composition, and annual brome species seeding to restore sites in the Northern Great Plains that composition. During the course of this study, both sites are dominated by annual brome species. At two contrast- experienced normal to above average annual precipita- ing sites, we compared two herbicide application rates tion compared to historical means (old-field, 1917–2011; that would facilitate the emergence and survival of seeded rangeland, 1972–2011) (Western Regional Climate Center species by decreasing annual brome species abundance. 2014; Figure 1). 148 • June 2015 ECOLOGICAL RESTORATION 33:2 Old-field Study Site Seeding treatments were applied the spring following the The old-field site (48°26'55.40" N, 109°51'57.45" W) was herbicide application (April 19, 2010 at the rangeland site; located at an elevation of 800 m, with fine loam soils and April 20, 2010 at the old-field sites). We used a 120 cm wide mean annual precipitation of 307 mm (Soil Survey Staff hoe seeder to sow eight rows in each main plot at 30 cm 2013, Western Regional Climate Center 2014). The site was spacing and two cm depth. We chose three species for each located in a former agricultural field which was plowed and location based on site characteristics (e.g., precipitation and seeded to a perennial grass mix in 2003 as part of the Con- soil type), fall-applied imazapic tolerance according to the servation Reserve Program (CRP). We initiated our study product label, and observations of local plant communities in 2009 and by then a mix of downy brome and Japanese (Anonymous 2008, Sheley et al. 2008). Species consisted of brome, hereafter referred to as brome, dominated the site, two grasses and a forb at each site, and seeding rates for with an associated substantial litter layer. When we started single species plots followed Sheley et al. (2008). Seeded our study, the existing non-brome plant community was species and rates at the old-field site were thickspike wheat- comprised of remnant perennial grasses seeded by the pri- grass (Elymus lanceolatus ‘Bannock’) at 12 kg pure live seed vate landowner in 2003 (western wheatgrass [Pascopyrum (PLS)/ha, pubescent wheatgrass (Thinopyrum intermedium smithii], green needlegrass [Stipa viridula], and crested ‘Luna’) at 16 kg PLS/ha, and purple prairieclover (Dalea wheatgrass [Agropyron cristatum]) and weedy perennial purpurea ‘Bismark’) at 9 kg PLS/ha. At the rangeland site, and annual forbs (dandelion [Taraxacum officinale], field we sowed western wheatgrass (‘Rosana’) at 18 kg PLS/ha, bindweed [Calystegia hederacea], yellow sweetclover [Meli- and pubescent wheatgrass and purple prairieclover at the lotus officinalis], prickly lettuce [Lactuca serriola], and same rates as above. Each species was sown singly, and in kochia [Bassia scoparia]). all combinations of two and three species for a total of eight seeding treatments including a non-seeded control. Rangeland Study Site For mixtures, we adjusted seeding rates so the total number The rangeland site (48°08'23.59" N, 107°52'15.58" W) was of seeds/m2 remained constant and species were seeded in located at an elevation of 725 m, with clay loam soils and equal proportions. mean annual precipitation of 329 mm (Soil Survey Staff Sampling was conducted in 2010 and 2011 to quantify 2013, Western Regional Climate Center 2014). This site was brome and plant community response to herbicide treat- on private land used primarily for cattle production and ments, and seeded species emergence (2010) and establish- had no history of plowing or cultivation.