Mountain roads shift native and non‐native plant species' ranges

Roads are known to act as corridors for dispersal of plant species. With their variable microclimate, role as corridors for species movement and reoccurring disturbance events, they show several characteristics that might influence range dynamics of both native and non‐native species. Previous research on plant species ranges in mountains however seldom included the effects of roads. To study how ranges of native and non‐native species differ between roads and adjacent vegetation, we used a global dataset of plant species composition along mountain roads. We compared average elevation and range width of species, and used generalized linear mixed models (GLMMs) to compile their range optimum and amplitude. We then explored differences between roadside and adjacent plots based on a species’ origin (native vs non‐native) and nitrogen and temperature affinity. Most non‐native species had on average higher elevational ranges and broader amplitudes in roadsides. Higher optima for non‐native species were associated with high nitrogen and temperature affinity. While lowland native species showed patterns comparable to those in non‐native species, highland native species had significantly lower elevational ranges in roadsides compared to the adjacent vegetation. We conclude that roadsides indeed change the elevational ranges of a variety of species. These changes are not limited to the expansion of non‐native species along mountain roads, but also include both upward and downward changes in ranges of native species. Roadsides may thus facilitate upward range shifts, for instance related to climate change, and they could serve as corridors to facilitate migration of alpine species between adjacent high‐elevation areas. We recommend including the effects of mountain roads in species distribution models to fine‐tune the predictions of range changes in a warming climate.     

Distribution and spread of environmental weeds along New?Zealand roadsides

Most non-native weeds and other naturalised plants are in the early stages of invasion into New Zealand landscapes. For this invasion to be controlled, even partially, it is important to understand the dominant routes, mechanisms, and rates of weed spread across landscapes. With their linear corridors of disturbed habitats, roadsides are thought to play a large role in the spread of some weeds. We used both new surveys and existing data to assess which of the 328 environmental weeds listed by the Department of Conservation are most frequently found and where on roadsides, and whether distribution patterns are consistent with linear dispersal. We also analysed historical survey data for relationships between reserve weediness and proximity to roads. We surveyed 340 plots of 100-m-long stretches of roadside across four regions and found between 2 and 19 environmental weeds per plot; 128 species in total (Chao estimate 148). Especially abundant were agricultural (weeds and cultivated) species, species that have been naturalised for well over 50 years, and species that disperse externally attached to vertebrates. While we purposefully sampled within 10 km of town limits, we found no strong effect of distance from town on roadside weed richness, including richness of just ornamentally sourced weeds. Instead, number of houses within 250?m and presence of an adjacent house or other residential structure were both important, as was presence of woody vegetation on and adjacent to roadsides. Reserves adjacent to roads had significantly higher weed richness than reserves further from roads, although the causal mechanisms are unclear. Our results suggest that while roadsides include suitable habitats for most environmental weeds, distributions are patchy and roads show little sign of acting as linear dispersal corridors, instead they largely reflect neighbouring land uses. As such, roadside weeds should best be managed as part of the wider landscape.     

The habitat and conduit functions of roads in the spread of three invasive plant species

Nonnative plant species commonly occur along roadsides, and populations are often assumed to invade by spread along the road axis. To distinguish between the function of roadsides as movement corridors and as habitat, nonnative plant species were surveyed along roads in deciduous forest sites in southeastern Ohio, USA. The importance of road proximity was tested by comparing nonnative species abundance in 100 m transects along roads with transects in undisturbed forest. Nonnative species were most abundant and most frequently observed in roadside sites in valleys. Three common species were chosen for closer scrutiny. In a seed sowing experiment roads and open sites proved to be better locations for the germination and growth of Microstegium vimineum than non-roadside and closed-canopy sites. Tussilago farfara and Rosa multiflora occurred in a small number of disjunct patches suggesting infrequent arrival in the sampled transects. Both species were strongly clustered at scales consistent with diffusive spread by vegetative growth and short-range seed dispersal. Comparisons of distributions parallel and perpendicular to roads showed no evidence for enhanced dispersal along the road axis. Microstegium distributions were correlated with local light availability implying site saturation. Microstegium micro-distributions suggested that spread along the road axis was facilitated by movement of dormant seeds in road maintenance. Thus, roadsides appear to function as both habitat and a conduit for population expansion, with the rate of spread dependent on the life history of the individual species. These results suggest a hierarchical process of regional invasion, with different dispersal mechanisms functioning at different spatial scales.     

Response of native and non-native ruderals to natural and human disturbance

The ruderal strategy is widely shared among non-native plants, providing a general explanation for the commonly observed positive effects of disturbance on invasions. How native ruderals respond to disturbance and how their abundance compares to that of non-native ruderals remains, however, poorly understood. Similarly, little is known about the role that disturbance type plays in the coexistence between native and non-native ruderals. We proposed that natural disturbance favors native over non-native ruderals, whereas novel anthropogenic disturbance favors non-natives over natives. To assess our general hypothesis, we conducted extensive field samplings in which we measured relative abundance, richness, and diversity of native and non-native ruderals in sites with natural and anthropogenic disturbance in central Argentina, a system where the ruderal strategy is common to a large number of native and non-native species. We found that natives dominated ruderal communities growing in recently burned grasslands, whereas non-natives dominated in roadsides. Additionally, the richness and diversity of native ruderal species were much greater than those of non-natives in sites with fire and in sites with grazing, but species richness and diversity did not differ between groups in roadsides. Because vegetation evolved with fire in our system and, in contrast, the construction and maintenance of roads is recent in it, these results support our hypothesis. Our work indicates that the ruderal strategy does not seem to suffice to explain why disturbance facilitates invasions. According to our data, species origin interacts with disturbance type to determine dominance in communities with coexisting native and non-native ruderals.     

Seed predation of non-native species along a precipitation gradient

Generalist seed predators are an important factor shaping non-native species invasion. Their effect is highly influenced by abiotic conditions, yet how the importance of this biotic filter changes in a gradient of abiotic conditions is still poorly understood. In this study, we assessed seed predation of non-native conifer species along a precipitation gradient in north-western Argentinean Patagonia. We performed a seed removal experiment over a 15-day period during the fall in sites where annual precipitation ranges from 600 to 1600 mm and vegetation ranges from steppe to forest. We used the three most common forestry species in the study area, including two species known to invade areas adjacent to plantations and one native species. Seed predation was higher in sites with higher precipitation, but no significant differences in seed predation were found among species. Small mammals were the dominant group predating seeds on this gradient. These findings highlight the importance of abiotic conditions mediating plant-granivore interactions, and the influence that abiotic conditions may have on biotic resistance to invasion.     

Settlements as a source for the spread of non-native plants into Central European suburban forests

Urbanization is considered as a major driver for biotic homogenization. Urbanization also promotes the dispersal of non-native species. This study examined the roles of suburban settlements and of the surrounding landscape composition for the spread of non-native plant species into adjacent mixed deciduous forests in Southern and Northwestern Switzerland. The number and abundance of native and non-native vascular plant species in both the ground vegetation and shrub layer were recorded in 15 forest sites situated adjacent to settlements and 15 control sites far from settlements. Various site and landscape characteristics were assessed in the surroundings (100 m radius) of the study sites. In both regions we found a higher number and larger abundance of non-native plant species in forest sites adjacent to settlements than in control forest sites. Furthermore, non-native plants were more frequently recorded close to roads and in sites surrounded by a large percentage cover of garden. All these effects were more pronounced in Southern Switzerland, a region with milder winter climate, than in Northwestern Switzerland. Our study showed that settlements are a source for the spread of non-native plant species into Central European suburban forests, and that the composition of the surrounding landscape matrix (e.g. traffic infrastructure, percentage cover of gardens) also affects the establishment of non-native plants.     

Effects of roads on alpine and subalpine plant species distribution along an altitudinal gradient on Mount Norikura, central Japan

We investigated the effects of roads on alpine and subalpine plant species distribution along an altitudinal gradient on Mount Norikura (3026 m a.s.l.), Japan. We examined the vegetation of herb and tree species shorter than 1.3 m along roadsides and adjacent natural vegetation at 200 m intervals between 1600 and 3000 m a.s.l. The timberline was at 2500 m a.s.l. Although the canopy opening was greater at the roadsides than in the natural vegetation, it was similar above the timberline. Soil cover and litter depth of the soil surface were less at roadsides than the natural vegetation, and gravel and rock cover were greater at roadsides. Species composition changed in similar directions from natural vegetation to roadsides along the altitudinal gradient. This direction was related to canopy opening and litter depth. Liliaceae, Ericaceae and Pinaceae were dominant families in the natural vegetation, and Asteraceae and Poaceae were greatest at the roadsides. Roadside plants were mostly herb species, while tree species increased in natural vegetation. Five exotic species were also observed at the roadsides. Sunny plant species gradually increased with altitude in the natural vegetation, indicated by the increase in canopy opening. By contrast, roadside plants were mostly sunny plant species irrespective of altitude. The number of lowland and montane species increased at the roadsides in the subalpine zone. Thus, roads strongly altered species composition of the natural vegetation along the altitudinal gradient probably because of changes in light and soil-surface conditions for growth and seedling establishment.     

Grassland connectivity by motor vehicles and grazing livestock

In addition to habitat loss and fragmentation, agricultural change has led to a change in seed dispersal processes in the rural landscape through a loss of structural and functional connectivity. Here, human‐mediated dispersal vectors are prevalent, and we explored whether the loss of connectivity via free‐ranging livestock could be mitigated by the increase in roads and motor vehicles. We found that structurally, 39% of all valuable semi‐natural grassland habitats in southern Sweden are adjacent to public road verges, which in the rural landscape are often considered to be suitable habitat for grassland species. Additionally, by collecting mud attached to cars and farming machinery and manure from livestock (cattle, horse, sheep) grazing semi‐natural grassland pasture, we found that motor vehicles are also capable seed dispersers. A similar number of species were dispersed by both vectors, although the composition of samples was quite different. Motor vehicles dispersed more grassland specialists than invasive species, although in much lower abundances than did grazing livestock. Despite these differences, motor vehicles were found to be able to disperse species with the same kinds of dispersal traits as livestock. A high number of seeds, species and specialists in manure samples means that greater movement of livestock is desirable to increase functional grassland connectivity. However, effective management could improve the suitability of roadsides as grassland corridors and increase the availability of seeds for long‐distance human‐mediated dispersal via cars and tractors. Our results suggest that in many rural landscapes, connectivity by road networks could help mediate habitat loss and fragmentation of grasslands. However, such effects can be context dependent, and the connectivity provided by roads could have serious negative consequences in other regions.     

Plant species-richness and invasion by exotics in relation to disturbance of wetland communities on the Riverine Plain,NSW

A combination of field surveys and seed-bank measurements were used to describe the macrophytic vegetation of intermittent wetlands on the Riverine Plain, in New South Wales. Three habitat types were sampled, representing wetland vegetation subjected to varying intensities of exogenous (human-induced) disturbance: swamps (least disturbed), roadside table drains (moderately disturbed), and rice crops (highly disturbed). Forty-two plant species were common to all three habitats and 36 species occurred in two of the three habitats. Of the 82 species that were restricted to a single habitat type, 33 were recorded for swamps, 19 for roadsides and 30 for rice fields. Species-richness (number of species per 50 m²) was found to be significantly lower in rice fields compared with either roadsides or swamps, which were not significantly different. Richness of native vascular species differed significantly among all habitat types, with swamps > roadsides > rice fields. Richness of exotic species also differed: roadsides > rice fields > swamps. Although the least disturbed habitat (swamps) supported the least number of exotic species, there was no further evidence of a positive relationship between degree of disturbance and degree of invasion. The results provided some support for the hypothesis that maximum species-richness occurs under conditions of moderate disturbance. A small negative correlation (r=?0.178, P < 0.01) was found between the number of exotic species and the number of native species occurring at a site. This suggests that, in this case, species-richness is not an important factor in the resistance of vegetation to invasion by exotic species. The vegetation of intermittent wetlands appears to have a relatively high degree of resistance to invasion. This is attributed to the high level of endogenous disturbance to which the native vegetation is adapted.     

Do positive interactions between marine invaders increase likelihood of invasion into natural and artificial habitats?

Positive species interactions such as facilitation are important for enabling species to persist, especially in stressful conditions, and the nature and strength of facilitation varies along physical and biological gradients. Expansion of coastal infrastructure is creating hotspots of invasive species which can spillover into natural habitats, but the role of positive species interactions associated with biological invasions remains understudied. Theory suggests that stronger biotic pressure in natural habitats inhibits invasion success. In space-limited marine systems, sessile organisms can overcome this limiting resource by settling as an epibiont on a substrate organism – basibiont. Using a series of spatially extensive surveys, we explored the role of invasive and native basibionts in providing habitat for other invasive and native epibionts, and tested whether environmental context (i.e. if the receiving habitat was natural or artificial), altered ecological outcomes. Overall, provision of space by basibionts was more important for invasive epibionts than for native epibionts but was dependent on the environmental context. Invasive basibionts facilitated invasive epibionts in natural habitats, and appeared to be more important for native epibionts in artificial habitats respectively. Native basibionts facilitated invasive, but not native epibionts in both natural and artificial habitats. These results advance our understanding of facilitation and highlight the idiosyncratic nature of biofouling and epibiosis, and the potentially important influence of environmental context. The degree to which native habitat-forming species versus invasive habitat-forming species either do or do not facilitate other native or non-native species is a rich area for investigation. Experimental work is required to disentangle the processes underpinning these patterns.     

Distribution of non-native invasive species and soil properties in proximity to paved roads and unpaved roads in a quartzitic mountainous grassland of southeastern Brazil (rupestrian fields)

One of the most important disturbances of roads is the facilitation of the increase of non-native invasive species into adjacent plant communities. The rupestrian fields of Serra do Cipó, a montane grassland ecosystem in southeastern Brazil, are recognized for their enormous richness of species and endemism rates. The presence of non-native invasive species in this ecosystem could threaten the existence of the native flora and its associated organisms. The aim of this study is to understand how non-native invasive species and native species are distributed along paved and unpaved roads, in a montaneous grassland ecosystem such as the Brazilian rupestrian fields. The two road surfaces provide differing gradients from their edges with respect to nutrients, soil chemical aspects and plant species diversity. High content of calcium at the roadside in the paved road resulted from the paving process, in which limestone gravel is used in one of the several paving phases. In these newly created habitats the toxicity of aluminum is drastically reduced and nutrient enriched, hence representing favorable sites from where non-native invasive species are capable to colonize and grow for undetermined period waiting the chance to invade the adjacent pristine habitats. Disturbances provoked by any natural or human-caused event can provide the opportunity for the non-native invasive species to colonize new plant communities.     

外来种对生物多样性的影响及其控制

外来种是那些借助自身力量或其它外界力量传播到其未曾分布过的地域 ,并且能进行繁殖传播的生物。外来种入侵已成为一种引人关注的现象。外来种通过竞争、捕食、牧食、改变生境和传播疾病等方式对本地生物产生威胁 ,影响本地生物多样性。外来种入侵成了生物多样性丧失的两个主要影响之一 (另一影响是生境的破坏 )。因此 ,探寻阻止或减少外来种入侵的方法很有必要。目前 ,人们运用机械法、化学法和生物控制法来控制外来种。在外来种入侵的初始阶段或外来种数量不多时 ,运用机械法较好。化学法有带来新环境污染的危险。生物控制法是应用天敌来防治或消灭有害生物。在外来种的控制上 ,生物控制法有一些成功的案例 ,但它并不是万能的。有些生物控制剂可能对非目标种产生影响 ,这要求在释放前对生物控制剂作更严格的检测。     

Bird communities of highway verges: Influence of adjacent habitat and roadside management

We have investigated the effects of landscape traversed and roadside structure on the use of highway verges by birds. Three contrasted landscapes were chosen in terms of human land use and vegetation structure: an intensive farmland, a pine plantation, and a matoral. The roadside sections varied in vegetation structure, width and profile. We recorded birds present in roadsides and adjacent habitats by transect counts over all seasons. Roadside bird species appeared for a great part similar to those of adjacent habitats. However, diversity and abundance in verges did not depend on that of adjacent habitats. Woody roadsides were comparable to hedges, as trees (and shrubs) in verges enhanced species richness and abundance of birds in the farmland and woodland sites. Width and profile of verges had less influence. In all sites, typical species of the habitat traversed partly avoided roadsides. On the contrary, numerous species associated with ‘rare’ habitats in one site preferred roadsides, provided that verge vegetation contrasted with the dominant habitat. It is concluded that birds responses to highways can vary greatly with landscape traversed and verge vegetation. Highway verges could be favorable to birds, if they constitute a complementary habitat to the dominant habitat within a landscape.     

In tallgrass prairie restorations,relatedness influences neighborhood-scale plant invasion while resource availability influences site-scale invasion

Attempting to control invasive plant species in tallgrass prairie restorations is time-consuming and costly, making improved approaches for predicting and reducing invasion imperative. Both biotic and abiotic factors mediate plant invasions, and can potentially be used by restoration managers to reduce invasion rates. Biotic factors such as plant species richness and phylogenetic diversity of the native community may impact invasion. Relatedness of invading species to those in recipient communities has also been shown to influence invasion success. However, the direction of this influence is variable, reflecting Darwin’s Naturalization Conundrum. Abiotic factors such as fire regime and soil factors may impact invasion by selecting against invasive species or indicating suitable habitats for them. We surveyed 17 tallgrass prairie restorations in Illinois, USA, to investigate the effects of biotic and abiotic factors on invasion by non-native plant species at two different scales. We predicted we would find support for Darwin’s Naturalization Hypothesis at the plot (neighborhood) scale with invasion by distantly related species, and find support for the Pre-adaptation Hypothesis at the site scale. We hypothesized that biotic factors would exert more influence at the neighborhood scale, while abiotic factors would be more influential at a coarser site scale. Contrary to our expectations, at the neighborhood scale we found that closely related invasive species are more likely to invade, supporting the Pre-adaptation Hypothesis. We found that native species richness and age of restoration were negatively correlated with invasion. At the site scale, soil organic matter [SOM] concentrations and heterogeneity in SOM were positively associated with the number of invasive species while pH heterogeneity was negatively associated. Restoration practitioners may be able to reduce plant invasions by increasing native species richness, and non-native species most closely related to the resident community should potentially be prioritized as those most likely to be highly invasive.     

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.     

Evidence that phylogenetically novel non-indigenous plants experience less herbivory

The degree to which biotic interactions influence invasion by non-indigenous species may be partly explained by the evolutionary relationship of these invaders with natives. Darwin’s naturalization hypothesis controversially proposes that non-native plants are more likely to invade if they lack close relatives in their new range. A possible mechanism for this pattern is that exotics that are more closely related to natives are more likely to share their herbivores, and thus will suffer more damage than phylogenetically isolated species. We tested this prediction using exotic plants in Ontario, Canada. We measured herbivore damage to 32 species of exotic plants in a common garden experiment, and 52 in natural populations. We estimated their phylogenetic distances from locally occurring natives in three ways: as mean distance (age) to all native plants, mean distance to native members of the same family, and distance to the closest native species. In the common garden, the proportion of leaves damaged and the average proportion of leaf area damaged declined with mean phylogenetic distance to native family relatives by late summer. Distance to native confamilials was a better predictor of damage than distance to the closest native species, while mean distance to the entire native plant community failed to predict damage. No significant patterns were detected for plants in natural populations, likely because uncontrolled site-to-site variation concealed these phylogenetic trends. To the extent that herbivory has negative demographic impacts, these results suggest that exotics that are more phylogenetically isolated from native confamilials should be more invasive; conversely, native communities should be more resistant to invasion if they harbor close familial relatives of potential invaders. However, the large scatter in this relationship suggests that these often are likely to be weak effects; as a result, these effects often may be difficult to detect in uncontrolled surveys of natural populations.     

Plant invasions along mountain roads: the altitudinal amplitude of alien Asteraceae forbs in their native and introduced ranges

Studying plant invasions along environmental gradients is a promising approach to dissect the relative importance of multiple interacting factors that affect the spread of a species in a new range. Along altitudinal gradients, factors such as propagule pressure, climatic conditions and biotic interactions change simultaneously across rather small geographic scales. Here we investigate the distribution of eight Asteraceae forbs along mountain roads in both their native and introduced ranges in the Valais (southern Swiss Alps) and the Wallowa Mountains (northeastern Oregon, USA). We hypothesised that a lack of adaptation and more limiting propagule pressure at higher altitudes in the new range restricts the altitudinal distribution of aliens relative to the native range. However, all but one of the species reached the same or even a higher altitude in the new range. Thus neither the need to adapt to changing climatic conditions nor lower propagule pressure at higher altitudes appears to have prevented the altitudinal spread of introduced populations. We found clear differences between regions in the relative occurrence of alien species in ruderal sites compared to roadsides, and in the degree of invasion away from the roadside, presumably reflecting differences in disturbance patterns between regions. Whilst the upper altitudinal limits of these plant invasions are apparently climatically constrained, factors such as anthropogenic disturbance and competition with native vegetation appear to have greater influence than changing climatic conditions on the distribution of these alien species along altitudinal gradients.     

The elephant in the room: the role of failed invasions in understanding invasion biology

Most species introductions are not expected to result in invasion, and species that are invasive in one area are frequently not invasive in others. However, cases of introduced organisms that failed to invade are reported in many instances as anecdotes or are simply ignored. In this analysis, we aimed to find common characteristics between non‐invasive populations of known invasive species and evaluated how the study of failed invasions can contribute to research on biological invasions. We found intraspecific variation in invasion success and several recurring explanations for why non‐native species fail to invade; these included low propagule pressure, abiotic resistance, biotic resistance, genetic constraints and mutualist release. Furthermore, we identified key research topics where ignoring failed invasions could produce misleading results; these include studies on historical factors associated with invasions, distribution models of invasive species, the effect of species traits on invasiveness, genetic effects, biotic resistance and habitat invasibility. In conclusion, we found failed invasions can provide fundamental information on the relative importance of factors determining invasions and might be a key component of several research topics. Therefore, our analysis suggests that more specific and detailed studies on invasion failures are necessary.     

Assessing the importance of disturbance, site conditions, and the biotic barrier for dandelion invasion in an Alpine habitat

Several factors have been identified as relevant in determining the abundance of non-native invasive species. Nevertheless, the relative importance of these factors will vary depending on the invaded habitat and the characteristics of the invasive species. Due to their harsh environmental conditions and remoteness, high-alpine habitats are often considered to be at low risk of plant invasion. However, an increasing number of reports have shown the presence and spread of non-native plant species in alpine habitats; thus, it is important to study which factors control the invasion process in these harsh habitats. In this study, we assessed the role of disturbance, soil characteristics, biotic resistance and seed rain in the establishment and abundance of the non-native invasive species Taraxacum officinale (dandelion) in the Andes of central Chile. By focusing on human-disturbed patches, naturally disturbed patches, and undisturbed patches, we did not find that disturbance per se, or its origin, affected the establishment and abundance of T. officinale. The abundance of this non-native invasive species was not negatively related to the diversity of native species at local scales, indicating no biotic resistance to invasion; instead, some positive relationships were found. Our results indicate that propagule pressure (assessed by the seed rain) and the abiotic soil characteristics are the main factors related to the abundance of this non-native invasive species. Hence, in contrast to what has been found for more benign habitats, disturbance and biotic resistance have little influence on the invasibility of T. officinale in this high-alpine habitat.     

High and dry or sunk and dunked: lessons for tallgrass prairies from quaking bogs

Northern Wisconsin bogs provide a natural experiment on butterfly population occurrence in a naturally highly fragmented vegetation type, which may provide insight on conserving butterflies in anthropogenically fragmented and degraded landscapes. We surveyed butterflies in bogs (about as unaffected by humans as possible, but naturally occurring over <1% of northern Wisconsin) primarily during 2002?C2009, with additional observations from 1986 to 2001. Different bog types had different bog-specialist butterfly faunas, but bog butterfly abundance also differed in similar vegetations among subregions. Some small isolated bogs held very high densities of specialist butterflies. Summer but not spring specialists frequented adjacent lowland roadsides and utilized a variety of non-native as well as native nectar sources. Paleo-entomology indicates that insects don??t evolve out of trouble; instead they move out of trouble. Given the low dispersal apparent today for species restricted to bogs, ??move?? might be better understood as ??hunkering?? within their vegetation as it expands and shrinks and moves around the landscape. Although bogs appeared to have more intact specialist butterfly faunas than tallgrass prairies (99.9% destroyed by human activities), bog butterflies do not live in average sites even in a relatively natural landscape. Just as bog butterflies are ??sunk and dunked?? in isolation, specialist butterflies elsewhere may have been left ??high and dry?? naturally, or are now due to human activities. Numerous studies have demonstrated that presence and abundance of specialist butterflies increase with increasing size and connectedness of habitat patches. But with long-term consistent vegetation, populations with high abundances in small isolated sites and with low numbers thinly occurring in large sites can be secure, as shown by bog butterflies.