Hansjörg Dietz

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.

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.

The role of bioclimatic origin, residence time and habitat context in shaping non-native plant distributions along an altitudinal gradient

An important factor influencing whether or not a non-native plant species becomes invasive is the climate in the area of introduction. To become naturalised in the new range, a species must either be climatically pre-adapted (climate matching), have a high phenotypic plasticity, or be able to adapt genetically, which in the latter case may take many generations. Furthermore, patterns of successful establishment across species might vary with habitat context. To address the interaction of these factors on non-native species richness, we recorded the presence of non-native annual plant species along an altitudinal gradient on Tenerife (Canary Islands, Spain). We compared the distributions of species differing in bioclimatic origin (Mediterranean and temperate) and time since introduction (old and recent introductions), and compared richness patterns of these groups in anthropogenic and natural habitats. Non-native species richness increased strongly from lowlands to mid-altitudes, but dropped sharply at the transition from anthropogenic to natural habitats, and thereafter declined with altitude in the natural habitat. This pattern indicates that the altitude effects reflected changes in both climate and habitat context. Mediterranean and temperate species were distributed similarly along the altitudinal gradient, and we found no effect of bioclimatic origin on species distributions. As almost all species present at the highest sites also occurred in the lowlands, we conclude that most species were introduced to lowland sites and were therefore pre-adapted to those climatic conditions (lowland introduction filter). The altitudinal ranges of species tended to increase with time since introduction, and the species reaching the highest altitudes were mostly old introductions. This effect of time was more pronounced among Mediterranean than temperate species. Thus, while climatic pre-adaptation is important for establishment along this altitudinal gradient, species tend to extend their altitudinal range with time.

Linear relationships between aboveground biomass and plant cover in low open herbaceous vegetation

Four herbaceous plant species of a sand dune area and several herb species of an open early-successional patch were investigated for the occurrence of a simple relationship between aboveground biomass and plant cover. Without exception linear regressions of aboveground biomass on plant cover were found with slope factors depending on the growth form of the species. These results suggest that (early) growth of herbaceous plants in low and (temporarily) open vegetation is not affected by possible constraints caused by a decreasing ratio of plant cover to aboveground biomass. The obtained linear relationships could be used for rapid non-destructive determination of aboveground biomass by image-analysis of cover data.

Determination of plant species cover by means of image analysis

. A method is described for the determination of plant species cover in herbaceous plant communities by means of image analysis. It is a computerized extension of the quadrat-charting method. The method was tested with four different computer systems for three different vegetation types, a shady lawn, a pioneer vegetation and a forest floor. We compared the results with vertical point-intercept analyses of the same stands. Image analysis revealed a high accuracy of the method ranging from < ± 1 to about ± 2% cover depending on the dominance of the species being analysed. The method is particularly suitable for the analysis of low herbaceous or unsaturated vegetation, dominance stands of broad-leaved plant species and for the analysis of cover changes in permanent plots.

Influence of light and nutrient conditions on seedling growth of native and invasive trees in the Seychelles

Several recent studies have shown that plant invasions can occur in resource-poor and relatively undisturbed habitats. It is, therefore, important to investigate whether and how life-history traits of species invasive in such habitats differ from those of species that are only invasive in disturbed and resource rich habitats. We compared the growth of seedlings of native and invasive tree species from nutrient-poor secondary forests in the tropical Seychelles. We hypothesised that the relative performance of the two groups would change predictably along resource gradients, with native species performing better at low levels of resource availability and invasive species performing better at higher levels. To test this hypothesis, we performed a common garden experiment using seedlings of six invasive and seven native tree species grown under three levels of light (65, 11 and 3.5% of ambient light) and two of nutrients (low and high). Due to large variation among species, differences in growth rates (RGR) were not significant among seedlings of the native and the invasive species. However, seedlings of the invasive species showed higher specific leaf areas (SLA) and higher leaf nutrient contents than seedlings of the native species. They also exhibited greater plasticity in biomass and nutrient allocation (i.e., greater plasticity in LAR, RSR and leaf nutrient contents) in response to varying resource availability. However, differences between the mean values of these parameters were generally small compared with variation within groups. We conclude that successful invaders on nutrient-poor soils in the Seychelles are either stress-tolerant, possessing growth traits similar to those of the native species, or fast-growing but adapted to nutrient-poor soils. In contrast, the more typical, fast-growing alien species with no particular adaptations to nutrient-poor soils seem to be restricted to relative nutrient-rich sites in the lowlands. The finding—that some introduced species thrive in resource-poor habitats—suggests that undisturbed habitats with low resource availability may be less resistant to plant invasions than was previously supposed.

Shifts in dominance of native and invasive plants in experimental patches of vegetation

Abstract While it is widely accepted that disturbance often promotes the process of plant invasion, few data are available on specific relationships between different types of disturbance and success of plant invaders. Nor is much known about variation in invader behaviour in response to changing composition of the dominant native species in the recipient habitats. In the middle Main valley near Wurzburg (Germany), two invasive Brassicaceae forbs, Bunias orientalis and Rorippa austriaca , are frequently associated with a small number of dominant native species in the herbaceous vegetation of productive, disturbed sites. Variation in the relative dominance of these species suggests high variability between sites in the factors influencing the local invasion process. In our study, we examined variation in regeneration mode, type and frequency of disturbance, and native species composition, and considered life history differences between the two invasive species as factors contributing to the distinct dominance patterns. These factors were tested in a controlled field experiment in which individuals of the two invasive species and four native species were grown in mixed stands for almost 3 years. The development of the plants was monitored by cover values and all plants were harvested at the end of the study. All factors investigated contributed considerably to the development of distinct dominance patterns in the species mixtures. Relative to the natives, both invasive species were promoted by regeneration from vegetative fragments as compared to development from sexual offspring. While both invasive species were generally promoted by disturbance relative to the natives, they differed in their response to distinct types of disturbance, and these differences were strongly affected by species composition. Our results indicate that invasion success of R. austriaca may depend more on soil disturbance, soil transport and deposition, while B. orientalis is expected to particularly expand at mown sites that do not have dense cover by meadow grasses. Whether the native resident vegetation is mainly composed of successional competitors that suffer from disturbance or of ruderal competitors that compete early and strongly with the invasive species may be of critical importance for invasion by the two species.

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.

Regeneration growth of the invasive clonal forb Rorippa austriaca (Brassicaceae) in relation to fertilization and interspecific competition

A special type of clonal growth, spread by lateral roots, ishypothesized to be a favourable trait of invasive, opportunistic plant speciesof disturbed habitats. We tested this hypothesis for the invasive forbRorippa austriaca (Brassicaceae). Regenerationfrom root fragments, subsequent vegetative spread and allocation patterns inrelation to varied nutrient supply and intensity and pattern of interspecificcompetition were analyzed in container experiments. Regeneration success fromroot fragments was 100% and clonal spread was rapid but vegetativeperformance was strongly reduced under unfertilized conditions and,particularly, when interspecific competition was present. While the ratio ofabove- to belowground bio-mass did not differ considerably betweentreatments, R. austriaca allocated a high amount ofresources to belowground growth resulting in low aboveground but highbelowground biomass at harvest time relative to the matrix vegetation.Differences in shoot number or biomass between simulated gaps and denselyvegetated quadrants in the containers were (relatively) weak.Reproductive effort was less reduced under low resource levels, and the clonesdid not set seed at all, irrespective of the treatment. Our results show thatclonal growth by lateral roots and plasticity in clonal growth patterns inR. austriaca promote both exploitation of gaps andnutrient-rich microsites and resistance to competitors. Such plasticity,combined with its ability to regenerate from widely-dispersed rootfragments, contribute to the ability of the species to invade and persistwithindisturbed and spatially heterogeneous habitats.