Tim Seipel

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.

Assembly of nonnative floras along elevational gradients explained by directional ecological filtering.

Nonnative species richness typically declines along environmental gradients such as elevation. It is usually assumed that this is because few invaders possess the necessary adaptations to succeed under extreme environmental conditions. Here, we show that nonnative plants reaching high elevations around the world are not highly specialized stress tolerators but species with broad climatic tolerances capable of growing across a wide elevational range. These results contrast with patterns for native species, and they can be explained by the unidirectional expansion of nonnative species from anthropogenic sources at low elevations and the progressive dropping out of species with narrow elevational amplitudes—a process that we call directional ecological filtering. Independent data confirm that climatic generalists have succeeded in colonizing the more extreme environments at higher elevations. These results suggest that invasion resistance is not conferred by extreme conditions at a particular site but determined by pathways of introduction of nonnative species. In the future, increased direct introduction of nonnative species with specialized ecophysiological adaptations to mountain environments could increase the risk of invasion. As well as providing a general explanation for gradients of nonnative species richness and the importance of traits such as phenotypic plasticity for many invasive species, the concept of directional ecological filtering is useful for understanding the initial assembly of some native floras at high elevations and latitudes.

Plant invasion at landscape and local scales along roadways in the mountainous region of the Greater Yellowstone Ecosystem

Roadways are increasingly recognized as common points of entry for non-native species into natural habitats in mountainous areas. Studies were conducted within the Greater Yellowstone Ecosystem from 2003 to 2007 to evaluate (1) landscape scale patterns of non-native plant richness along roadways, and (2) local scale factors influencing native and non-native plant richness and cover, and surrogate non-native plant (SNP) emergence in an invaded habitat. At the landscape scale, non-native plant richness decreased with increased elevation and increased distance from the road, and was positively correlated to the proportion of plots with signs of disturbance. Non-native plant richness also varied by habitat type: sagebrush steppe had the highest and alpine the lowest. At the local scale, in sagebrush steppe, SNP emergence was negatively associated with increased distance from the road, and percent cover of litter was positively associated with SNP emergence. The proportion of non-native plant cover and richness decreased, while the proportion of native cover and native species richness increased with distance from road. Our study suggests that landscape scale variables such as elevation and habitat type influence non-native plant success, and that while local conditions adjacent to the road may be favorable for non-native plants, factors which vary at the local scale can also effect non-native plant establishment away from the roadside. This highlights the need for studies to evaluate multiple scales when assessing patterns and processes driving non-native plant invasions, and suggests that sagebrush steppe may be resistant to invasion as long as it remains undisturbed.

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.

Will climate change increase the risk of plant invasions into mountains

Mountain ecosystems have been less adversely affected by invasions of non-native plants than most other ecosystems, partially because most invasive plants in the lowlands are limited by climate and cannot grow under harsher high-elevation conditions. However, with ongoing climate change, invasive species may rapidly move upwards and threaten mid-, and then high-elevation mountain ecosystems. We evaluated this threat by modeling the current and future habitat suitability for 48 invasive plant species in Switzerland and New South Wales, Australia. Both regions had contrasting climate interactions with elevation, resulting in possible different responses of species distributions to climate change. Using a species distribution modeling approach that combines data from two spatial scales, we built high-resolution species distribution models (≤ 250 m) that account for the global climatic niche of species and also finer variables depicting local climate and disturbances. We found that different environmental drivers limit the elevation range of invasive species in each of the two regions, leading to region-specific species responses to climate change. The optimal suitability for plant invaders is predicted to markedly shift from the lowland to the montane or subalpine zone in Switzerland, whereas the upward shift is far less pronounced in New South Wales where montane and subalpine elevations are already suitable. The results suggest that species most likely to invade high elevations in Switzerland will be cold-tolerant, whereas species with an affinity to moist soils are most likely to invade higher elevations in Australia. Other plant traits were only marginally associated with elevation limits. These results demonstrate that a more systematic consideration of future distributions of invasive species is required in conservation plans of not yet invaded mountainous ecosystems.

Plant invasions into mountains and alpine ecosystems: current status and future challenges

Recent years have seen a surge of interest in understanding patterns and processes of plant invasions into mountains. Here, we synthesise current knowledge about the spread of non-native plants along elevation gradients, emphasising the current status and impacts that these species have in alpine ecosystems. Globally, invasions along elevation gradients are influenced by propagule availability, environmental constraints on population growth, evolutionary change and biotic interactions. The highest elevations are so far relatively free from non-native plants. Nonetheless, in total nearly 200 non-native plant species have been recorded from alpine environments around the world. However, we identified only three species as specifically cold-adapted, with the overwhelming majority having their centres of distribution under warmer environments, and few have substantial impacts on native communities. A combination of low propagule availability and low invasibility likely explain why alpine environments host few non-native plants relative to lowland ecosystems. However, experiences in some areas demonstrate that alpine ecosystems are not inherently resistant to invasions. Furthermore, they will face increasing pressure from the introduction of pre-adapted species, climate change, and the range expansion of native species, which are already causing concern in some areas. Nonetheless, because they are still relatively free from non-native plants, preventative action could be an effective way to limit future impacts of invasions in alpine environments.

Farming system and wheat cultivar affect infestation of, and parasitism on, Cephus cinctus in the Northern Great Plains: Cephus cinctus infestation on wheat cultivars

BACKGROUND Cephus cinctus infestation causes

Temperature and alternative hosts influence Aceria tosichella infestation and Wheat streak mosaic virus infection

350 million in annual losses in the Northern Great Plains. We compared infestation and parasitism of C. cinctus in spring (including Kamut; Triticum turgidum ssp. turanicum) and winter wheat cultivars grown in organic and conventional fields in Montana, USA. In the greenhouse, we compared C. cinctus preference and survival in Kamut, Gunnison, and Reeder spring wheat cultivars. RESULTS Stems cut by C. cinctus varied by farming system and the seasonality of the wheat crop. No stems of Kamut in organic fields were cut by C. cinctus, but 1.5% [±0.35% standard error (SE)] of stems in conventional spring wheat, 5% (±0.70% SE) of stems in organic winter wheat, and 20% (±0.93% SE) of stems in conventional winter wheat fields were cut by C. cinctus. More larvae of C. cinctus were parasitized in organic (27 ± 0.03% SE) compared with conventional (5 ± 0.01% SE) winter wheat fields. Cephus cinctus oviposition, survival, and the number of stems cut were lowest in Kamut compared with Gunnison and Reeder. CONCLUSION Cephus cinctus infestation was more common in winter wheat than in spring wheat. Organic fields with fewer cut stems also supported more parasitoids. Kamut is a genetic resource for developing C. cinctus-resistant cultivars.

Running off the road: roadside non-native plants invading mountain vegetation

Wheat streak mosaic, caused by Wheat streak mosaic virus (WSMV; family Potyviridae), is the most important and common viral disease of wheat (Triticum aestivum L.) in the Great Plains of North America. WSMV is transmitted by the wheat curl mite (WCM; Aceria tosichella). We evaluated how mean daily temperatures, cumulative growing degree-days, day of the year, and surrounding alternative host identity affected WCM infestation and WSMV infection of wheat from late summer through early autumn in Montana, United States. Cumulative growing degree-days, warm mean daily temperatures (i.e., >10°C), and surrounding alternative hosts interacted to alter risk of WCM infestation and WSMV infection. Wheat surrounded by Bromus tectorum L. and preharvest volunteer wheat had WCM infestation and WSMV infection rates of 88% in years when the mean daily temperature was 15°C in October, compared with 23% when surrounded by bare ground, and <1% when the temperature was 0°C regardless of surrounding alternative host. Mean daily temperatures in the cereal-growing regions of Montana during autumn are marginally conducive to WCM population growth and movement. As the region continues to warm, the period of WCM movement will become longer, potentially increasing the frequency of WSMV outbreaks.

Processes at multiple scales affect richness and similarity of non-native plant species in mountains around the world

Prevention is regarded as a cost-effective management action to avoid unwanted impacts of non-native species. However, targeted prevention can be difficult if little is known about the traits of successfully invading non-native species or habitat characteristics that make native vegetation more resistant to invasion. Here, we surveyed mountain roads in seven regions worldwide, to investigate whether different species traits are beneficial during primary invasion (i.e. spread of non-native species along roadside dispersal corridors) and secondary invasion (i.e. percolation from roadsides into natural adjacent vegetation), and to determine if particular habitat characteristics increase biotic resistance to invasion. We found primary invasion up mountain roads tends to be by longer lived, non-ruderal species without seed dispersal traits. For secondary invasion, we demonstrate that both traits of the non-native species and attributes of the receiving natural vegetation contribute to the extent of invasion. Non-native species that invade natural adjacent vegetation tend to be shade and moisture tolerant. Furthermore, non-native species invasion was greater when the receiving vegetation was similarly rich in native species. Our results show how mountain roads define which non-native species are successful; first by favouring certain traits in mountain roadsides (the key dispersal pathway to the top), and secondly by requiring a different set of traits when species invade the natural adjacent vegetation. While patterns in species traits were observed at a global level, regional abiotic and biotic variables largely generated region-specific levels of response, suggesting that management should be regionally driven.