Catherine G. Parks

Ain't no mountain high enough: plant invasions reaching new elevations

Most studies of invasive species have been in highly modified, lowland environments, with comparatively little attention directed to less disturbed, high-elevation environments. However, increasing evidence indicates that plant invasions do occur in these environments, which often have high conservation value and provide important ecosystem services. Over a thousand non-native species have become established in natural areas at high elevations worldwide, and although many of these are not invasive, some may pose a considerable threat to native mountain ecosystems. Here, we discuss four main drivers that shape plant invasions into high-elevation habitats: (1) the (pre-)adaptation of non-native species to abiotic conditions, (2) natural and anthropogenic disturbances, (3) biotic resistance of the established communities, and (4) propagule pressure. We propose a comprehensive research agenda for tackling the problem of plant invasions into mountain ecosystems, including documentation of mountain invasion patterns at multiple scales, experimental studies, and an assessment of the impacts of non-native species in these systems. The threat posed to high-elevation biodiversity by invasive plant species is likely to increase because of globalization and climate change. However, the higher mountains harbor ecosystems where invasion by non-native species has scarcely begun, and where science and management have the opportunity to respond in time.

Establishment of parallel altitudinal clines in traits of native and introduced forbs

Due to altered ecological and evolutionary contexts, we might expect the responses of alien plants to environmental gradients, as revealed through patterns of trait variation, to differ from those of the same species in their native range. In particular, the spread of alien plant species along such gradients might be limited by their ability to establish clinal patterns of trait variation. We investigated trends in growth and reproductive traits in natural populations of eight invasive Asteraceae forbs along altitudinal gradients in their native and introduced ranges (Valais, Switzerland, and Wallowa Mountains, Oregon, USA). Plants showed similar responses to altitude in both ranges, being generally smaller and having fewer inflorescences but larger seeds at higher altitudes. However, these trends were modified by region-specific effects that were independent of species status (native or introduced), suggesting that any differential performance of alien species in the introduced range cannot be interpreted without a fully reciprocal approach to test the basis of these differences. Furthermore, we found differences in patterns of resource allocation to capitula among species in the native and the introduced areas. These suggest that the mechanisms underlying trait variation, for example, increasing seed size with altitude, might differ between ranges. The rapid establishment of clinal patterns of trait variation in the new range indicates that the need to respond to altitudinal gradients, possibly by local adaptation, has not limited the ability of these species to invade mountain regions. Studies are now needed to test the underlying mechanisms of altitudinal clines in traits of alien species.

Plant Invasions in Mountains: Global Lessons for Better Management

Abstract Mountains are one of few ecosystems little affected by plant invasions. However, the threat of invasion is likely to increase because of climate change, greater anthropogenic land use, and continuing novel introductions. Preventive management, therefore, will be crucial but can be difficult to promote when more pressing problems are unresolved and predictions are uncertain. In this essay, we use management case studies from 7 mountain regions to identify common lessons for effective preventive action. The degree of plant invasion in mountains was variable in the 7 regions as was the response to invasion, which ranged from lack of awareness by land managers of the potential impact in Chile and Kashmir to well-organized programs of prevention and containment in the United States (Hawaii and the Pacific Northwest), including prevention at low altitude. In Australia, awareness of the threat grew only after disruptive invasions. In South Africa, the economic benefits of removing alien plants are well recognized and funded in the form of employment programs. In the European Alps, there is little need for active management because no invasive species pose an immediate threat. From these case studies, we identify lessons for management of plant invasions in mountain ecosystems: (i) prevention is especially important in mountains because of their rugged terrain, where invasions can quickly become unmanageable; (ii) networks at local to global levels can assist with awareness raising and better prioritization of management actions; (iii) the economic importance of management should be identified and articulated; (iv) public acceptance of management programs will make them more effective; and (v) climate change needs to be considered. We suggest that comparisons of local case studies, such as those we have presented, have a pivotal place in the proactive solution of global change issues.

Modeling Tamarisk (Tamarix spp.) Habitat and Climate Change Effects in the Northwestern United States

Abstract Tamarisk species are shrubs or small trees considered by some to be among the most aggressively invasive and potentially detrimental exotic plants in the United States. Although extensively studied in the southern and interior west, northwestern (Oregon, Washington, and Idaho) distribution and habitat information for tamarisk is either limited or lacking. We obtained distribution data for the northwest, developed a habitat suitability map, and projected changes in habitat due to climate change in a smaller case study area using downscaled climate data. Results show extensive populations of tamarisk east of the Cascade Mountains. Despite the perceived novelty of tamarisk in the region, naturalized populations were present by the 1920s. Major population centers are limited to the warmest and driest environments in the central Snake River Plain, Columbia Plateau, and Northern Basin and Range. Habitat suitability model results indicate that 21% of the region supports suitable tamarisk habitat. Less than 1% of these areas are occupied by tamarisk; the remainder is highly vulnerable to invasion. Although considerable uncertainty exists regarding future climate change, we project a 2- to 10-fold increase in highly suitable tamarisk habitat by the end of the century. Our habitat suitability maps can be used in “what if” exercises as part of planning, detection, restoration, management, and eradication purposes. Nomenclature: Tamarisk, species in the genus Tamarix L., primarily Tamarix chinensis Lour. and Tamarix ramosissima Ledeb. and their hybrids.

Plant Invasions into Mountain Protected Areas: Assessment, Prevention and Control at Multiple Spatial Scales

Mountains are of great significance for people and biodiversity. Although often considered to be at low risk from alien plants, recent studies suggest that mountain ecosystems are not inherently more resistant to invasion than other types of ecosystems. Future invasion risks are likely to increase greatly, in particular due to climate warming and increased human land use (e.g. intensification of human activities, human population growth, and expansion of tourism). However, these risks can be reduced by minimising anthropogenic disturbance in and around protected areas, and by preventing the introduction of potentially invasive alien plants into these areas, particularly at high elevations. Sharing information and experiences gained in different mountainous areas is important for devising effective management strategies. We review current knowledge about plant invasions into mountains, assembling evidence from all continents and across different climate zones, and describe experiences at local to global scales in preventing and managing plant invasions into mountain protected areas. Our findings and recommendations are also relevant for managing native species that expand to higher elevations.

Age structure and age-related performance of sulfur cinquefoil (Potentilla recta)

Abstract Age distributions of sulfur cinquefoil populations were determined on sites that were historically grazed, cultivated, and mechanically disturbed. From 12 sites, a total of 279 reproductively active plants were collected and aged by using herbchronology (counting rings in the secondary root xylem of the root crown) to (1) estimate the age structure of the populations, (2) relate plant size and flower production to plant age, and (3) examine the relation of population age structure to environmental variables and disturbance history. Results indicated that the mean age for all sampled plants was 3.5 (± 1.74 SD) yr and ranged from 1 to 10 yr. Age was not related to number of flowers, plant size (number of stems per plant or plant height), or site disturbance type but was positively correlated with site elevation (P < 0.001). The pooled age distribution from all 12 sites was right-skewed with fewer old plants than young plants. We conclude that sulfur cinquefoil plants sampled in northeast Oregon are able to colonize, establish, and reproduce at disturbed sites rapidly. We suggest that herbchronology may be a useful technique to improve understanding of invasion biology and ecology for invasive plant species that form annual rings. Nomenclature: Sulfur cinquefoil, Potentilla recta L. PTLRC.

Multiscale Detection of Sulfur Cinquefoil Using Aerial Photography

Abstract We evaluated the effectiveness of natural color aerial photography as a tool to improve detection, monitoring, and mapping of sulfur cinquefoil (Potentilla recta L.) infestations. Sulfur cinquefoil is an exotic perennial plant invading interior Pacific Northwest rangelands. Because sulfur cinquefoil produces distinctive pale yellow flowers, we timed aerial photography for early July, when the plant was at peak bloom. Photography was collected at 3 spatial scales (1:3 000, 1:6 000, and 1:12 000). A grid with 250-m spacing was superimposed over photographs of the entire study area using geographic information systems. At each grid intersection point (n = 80), we visually analyzed the photographs within a 404.7-m2 (0.1 acre) circular plot, recorded sulfur cinquefoil presence, and estimated sulfur cinquefoil percent cover. Sample points on the grid were then located in the field using a global positioning system. Field data collected at each point included sulfur cinquefoil presence, percent cover, and stem density; and total vegetation composition and percent cover by life form. Results indicate that the accuracy of detecting sulfur cinquefoil increased from small to large scale. At the 1:3 000 scale, sulfur cinquefoil presence was correctly identified in 76.9% of the sites, whereas at the 1:6 000 and 1:12 000 scales, infestations were identified in 67.9% and 59.1% of the sites, respectively. Low-density infestations (< 1% cover) were detected at all scales. Accuracy of percent cover estimates ranged from 33.8% to 38.0% across scales. Although tree canopy hindered detection, our results indicate that aerial photography can be used to detect sulfur cinquefoil infestations in open forests and rangelands in the Intermountain West.

Seed Production and Dispersal of Sulfur Cinquefoil in Northeast Oregon

Abstract Sulfur cinquefoil (family Rosaceae) is an invasive, herbaceous perennial, native to Eurasia. It has wide ecological amplitude and has become established throughout North America in numerous habitat types. Sulfur cinquefoil reproduces only by seed (achenes); however, little is known about its regenerative strategy or reproductive biology. To improve understanding of the mechanisms of expansion for sulfur cinquefoil, we quantified seed production and measured seed dispersal at sites infested with sulfur cinquefoil in different habitats in northeast Oregon. Seed dispersal was measured by using sticky traps (30 × 100 cm, replaced every 2 weeks) radiating in 4 cardinal directions from individual source plants. Estimated seed production for 2 years (2001 and 2002) was nearly 4 times higher than previously reported (≈6 000 seeds per plant; range ≈ 2 620–15 150 seeds per plant). For most sites, seed production was similar in both years. However, site, year, and their interaction (site × year) had significant influence on flower and stem production. Seeds were dispersed from July through mid-October 2001, although almost 40% of the seeds were captured between mid-July and mid-August. Dispersal followed a classic decay function; approximately 83% of the seeds were captured within 60 cm of the source plants. Once sulfur cinquefoil reaches a site, it appears to spread and persist by releasing numerous seeds near the parent plants, thereby forming increasingly dense stands.

Landscape Factors Influencing the Abundance and Dominance of the Invasive Plant Potentilla Recta

Abstract Little is known about the relative importance of environmental, biotic, historical, and spatial factors that influence invasive plant abundance, dominance, and distribution across landscapes. We identified factors that influence the abundance and dominance of Potentilla recta L. (sulfur cinquefoil) in bunchgrass grasslands of northeastern Oregon to better understand the conditions under which it becomes a major component of plant communities. We estimated P. recta stem density and dominance from field measurements across the landscape and used classification and regression tree analyses to assess the importance of environmental, biotic, spatial, and historical factors in explaining P. recta presence, stem density, and dominance. Plots were sampled within a systematic grid with 250-m spacing within our 6.5-km2 study landscape. At each sample point we recorded P. recta presence, stem density, and dominance as well as 11 biological, environmental, spatial, and historical variables. P. recta was widely distributed, with stem densities in occupied plots averaging 5.8 stems • m−2 and dominance values ranging from 1% to 52%. Percent cover of bare ground was the most important variable to predict the presence of P. recta, though the model fit was poor, likely because the entire study area is suitable for P. recta establishment. A strong relationship between P. recta dominance and habitat type (r2  =  67.5%) was found, with dominance greatest in old fields on relatively flat slopes (mean dominance of 34.1%). Dominance estimates were ≤ 1% in plots located in forest, shrub, and grassland habitats. Factors that make old fields susceptible to dominance remain unknown, though microsite conditions that increase P. recta seedling survival rates and limited native propagule availability due to previous cultivation may be involved. Since old fields are found throughout the region, are highly susceptible to P. recta invasion, and represent a source of seeds, containment and restoration activities should focus on these areas.

Non-Native Plant Invasion along Elevation and Canopy Closure Gradients in a Middle Rocky Mountain Ecosystem

Mountain environments are currently among the ecosystems least invaded by non-native species; however, mountains are increasingly under threat of non-native plant invasion. The slow pace of exotic plant invasions in mountain ecosystems is likely due to a combination of low anthropogenic disturbances, low propagule supply, and extreme/steep environmental gradients. The importance of any one of these factors is debated and likely ecosystem dependent. We evaluated the importance of various correlates of plant invasions in the Wallowa Mountain Range of northeastern Oregon and explored whether non-native species distributions differed from native species along an elevation gradient. Vascular plant communities were sampled in summer 2012 along three mountain roads. Transects (n = 20) were evenly stratified by elevation (~70 m intervals) along each road. Vascular plant species abundances and environmental parameters were measured. We used indicator species analysis to identify habitat affinities for non-native species. Plots were ordinated in species space, joint plots and non-parametric multiplicative regression were used to relate species and community variation to environmental variables. Non-native species richness decreased continuously with increasing elevation. In contrast, native species richness displayed a unimodal distribution with maximum richness occurring at mid–elevations. Species composition was strongly related to elevation and canopy openness. Overlays of trait and environmental factors onto non-metric multidimensional ordinations identified the montane-subalpine community transition and over-story canopy closure exceeding 60% as potential barriers to non-native species establishment. Unlike native species, non-native species showed little evidence for high-elevation or closed-canopy specialization. These data suggest that non-native plants currently found in the Wallowa Mountains are dependent on open canopies and disturbance for establishment in low and mid elevations. Current management objectives including restoration to more open canopies in dry Rocky Mountain forests, may increase immigration pressure of non-native plants from lower elevations into the montane and subalpine zones.