Think globally, measure locally: The MIREN standardized protocol for monitoring plant species distributions along elevation gradients.

Climate change and other global change drivers threaten plant diversity in mountains worldwide. A widely documented response to such environmental modifications is for plant species to change their elevational ranges. Range shifts are often idiosyncratic and difficult to generalize, partly due to variation in sampling methods. There is thus a need for a standardized monitoring strategy that can be applied across mountain regions to assess distribution changes and community turnover of native and non-native plant species over space and time. Here, we present a conceptually intuitive and standardized protocol developed by the Mountain Invasion Research Network (MIREN) to systematically quantify global patterns of native and non-native species distributions along elevation gradients and shifts arising from interactive effects of climate change and human disturbance. Usually repeated every five years, surveys consist of 20 sample sites located at equal elevation increments along three replicate roads per sampling region. At each site, three plots extend from the side of a mountain road into surrounding natural vegetation. The protocol has been successfully used in 18 regions worldwide from 2007 to present. Analyses of one point in time already generated some salient results, and revealed region-specific elevational patterns of native plant species richness, but a globally consistent elevational decline in non-native species richness. Non-native plants were also more abundant directly adjacent to road edges, suggesting that disturbed roadsides serve as a vector for invasions into mountains. From the upcoming analyses of time series, even more exciting results can be expected, especially about range shifts. Implementing the protocol in more mountain regions globally would help to generate a more complete picture of how global change alters species distributions. This would inform conservation policy in mountain ecosystems, where some conservation policies remain poorly implemented.

Establishment of parallel altitudinal dines 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.

Effects of landscape structure on movement patterns of the flightless bush cricket Pholidoptera griseoaptera.

Because the viability of a population may depend on whether individuals can disperse, it is important for conservation planning to understand how landscape structure affects movement behavior. Some species occur in a wide range of landscapes differing greatly in structure, and the question arises of whether these species are particularly versatile in their dispersal or whether they are composed of genetically distinct populations adapted to contrasting landscapes. We performed a capture-mark-resight experiment to study movement patterns of the flightless bush cricket Pholidoptera griseoaptera (De Geer 1773) in two contrasting agricultural landscapes in France and Switzerland. The mean daily movement of P. griseoaptera was significantly higher in the landscape with patchily distributed habitat (Switzerland) than in the landscape with greater habitat connectivity (France). Net displacement rate did not differ between the two landscapes, which we attributed to the presence of more linear elements in the connected landscape, resulting in a more directed pattern of movement by P. griseoaptera. Significant differences in the movement patterns between landscapes with contrasting structure suggest important effects of landscape structure on movement and dispersal success. The possibility of varying dispersal ability within the same species needs to be studied in more detail because this may provide important information for sustainable landscape planning aimed at maintaining viable metapopulations, especially in formerly well-connected landscapes.

Recognition that causal processes change during plant invasion helps explain conflicts in evidence.

Despite intensive research, we still have no general understanding of why plant invasions occur. Many different mechanisms of plant invasions have been proposed, but studies designed to investigate them often produce inconsistent results. It remains unclear whether this unsatisfying state of affairs reflects the complexity of the real world (in which every invasion is unique) or the failure to identify the key processes driving most plant invasions. Here we argue that greater generalization is possible, but only if we recognize that the ecological and evolutionary processes enabling a species to advance into a new area change during the course of an invasion. In our view, an invasion can often usefully be subdivided into a primary phase, in which the abundance of an often preadapted species increases rapidly (typically in resource-rich, disturbed habitats), and a secondary phase, in which further spread is contingent upon plastic responses or genetic adaptation to new ecological circumstances. We present various examples to show how this partitioning of the invasion phase sensu stricto produces new hypotheses about the processes underlying plant invasions. Some of these hypotheses can be conveniently tested by investigating plant invasions along strong environmental gradients such as those that occur in mountainous regions.

Evidence for life history changes in high-altitude populations of three perennial forbs.

Relatively little is known about how the life histories of perennial forb species, and especially their lifetime patterns of growth, vary across environmental gradients. We used a post hoc approach (herb-chronology) to determine plant age and previous growth (width of successive annual rings in roots) in three species of perennial forb (two long-lived species [Penstemon venustus, Lupinus laxiflorus] and one short-lived [Rudbeckia occidentalis]) along a 1000-m altitudinal gradient in the Wallowa Mountains (northeast Oregon, USA). Plants from the highest altitude tended to be considerably older and produced up to five times as many flowering shoots as lowland plants. In addition, mean ring widths of high-altitude plants were about half those of lowland plants. In plants from low and intermediate altitudes, ring width either decreased linearly or varied inconsistently during the life of the plant. In contrast, ring widths of high-altitude plants increased at first and later decreased, resulting in curvilinear growth trajectories that were highly consistent among species. Together, these data for three ecologically distinct forb species provide evidence of a consistent shift toward more conservative and strongly constrained life histories at higher altitudes. More generally, the results indicate the possible importance of changes in selection pressures across strong environmental gradients on life history strategies within a single species.

Population age structure and reproductive behavior of the monocarpic perennial Heracleum mantegazzianum (Apiaceae) in its native and invaded distribution ranges.

Many invasive species are benign in their native region-are there interactions between their key traits and the new habitats that explain invasion success? The giant perennial herb Heracleum mantegazzianum is a problematic invader in Europe and is also naturalized in North America. We compared its population structure and reproductive behavior in the native (W. Caucasus) and invaded (Czech Republic) areas in managed (pastures) and unmanaged sites. The age structure of the populations and age at flowering were analyzed using herb-chronology, a method based on counting annual rings in the secondary xylem of roots. The species was strictly monocarpic; most plants in unmanaged sites in the invaded range flowered in the third and fourth yr (maximum 12 yr). In unmanaged habitats, plants from the native range flowered later than those from the invaded range. In both ranges, flowering was delayed in managed sites where the population density was higher and most plants flowered around the fifth year. Reproductive output of individual plants was neither related to population density nor to age at flowering. More favorable climatic conditions in the invaded region, together with increased chances for dispersal in a densely colonized central Europe, seemed to allow the massive invasion.

Demographic and genetic invasion history of a 9-year-old roadside population of Bunias orientalis L. (Brassicaceae).

The population history of a 9-year-old roadside population of the invasive plant Bunias orientalis was reconstructed by demographic analysis including size, position, age (determined by herbchronology) and RAPD-PCR patterns of individual plants. We evaluated emerging patterns of population growth and genetic structure during a full period of population development under typical site conditions (anthropogenic disturbance) and their possible consequences for the invasion potential of the species. The population has grown rapidly and continuously (though with slowing geometric population increase) during the 9 years since its foundation, filling the space available in the study area. Genetic variation (RAPD markers) was already high in the founder cohorts and remained at the same level throughout population development (variance fluctuations <15%). Both results may be related to the mowing management at the site which seems to promote population growth of B. orientalis relative to other co-occuring species and to prevent the genetic drift and the development of spatial genetic structure that would be expected under isolation-by-distance models. Large founder plants had comparatively low genetic variance and were more closely related to younger cohorts than were small founder plants, indicating that selection acted during population development. Overall, the current anthropogenic disturbance regimes may contribute to high genetic variability by artificially increasing gene flow and thereby promoting the adaptability of invasive species to the often unpredictable conditions at disturbed sites. Our approach using retrospective demographic investigation allows the detection of spatio-temporal microscale patterns in genetic and phenotypic variation. Thus it allows a thorough understanding of local invasions of perennial herbaceous plants.