Biotic constraints on the invasion of diffuse knapweed (Centaurea diffusa) in North American grasslands

Knapweeds (Centaurea spp.) are among the most invasive of non-indigenous plant species that have colonized western North America over the last century. We conducted a 4-year experiment in a reconstructed grassland to test hypotheses related to the ability of grasslands to resist the invasion of diffuse knapweed (C. diffusa). We experimentally invaded C. diffusa and three native species into areas where we manipulated soil nitrogen (N) and phosphorus (P) availability and removed extant grasses to reduce competition. We evaluated the growth response of these species to these resources and competitive manipulations. Of the native species that were experimentally added, only one species, Ratibida pinnata (prairie coneflower), established in any numbers. Establishment values in intact vegetation were low for both species, but establishment by C. diffusa (0.02%) clearly outperformed that of R. pinnata (0.001%). Under reduced grass competition, establishment was enhanced, but the values for C. diffusa (0.68%) were not statistically different from those of R. pinnata (0.57%). Neither species performed better under higher soil nutrients in the presence of competing grasses. In plots with both species, biomass of the two planted species was positively correlated, but the biomass of both species was negatively correlated with non-added weedy species. Subsequent harvests of C. diffusa indicated that establishment was enhanced in treatments with higher soil nutrients but that the biomass of these plants could only be enhanced when plant competition was also reduced. These results indicate that C. diffusa can establish in intact grasslands at rates higher than natives, but opportunism rather than competitive ability best describes the invasiveness of C. diffusa. Thus, the mechanisms contributing to the establishment of this knapweed species are different from factors identified as contributing to the dominance of this invader.     

Evaluating Ecosystem Services Provided by Non-Native Species: An Experimental Test in California Grasslands

The concept of ecosystem services – the benefits that nature provides to human''s society – has gained increasing attention over the past decade. Increasing global abiotic and biotic change, including species invasions, is threatening the secure delivery of these ecosystem services. Efficient evaluation methods of ecosystem services are urgently needed to improve our ability to determine management strategies and restoration goals in face of these new emerging ecosystems. Considering a range of multiple ecosystem functions may be a useful way to determine such strategies. We tested this framework experimentally in California grasslands, where large shifts in species composition have occurred since the late 1700''s. We compared a suite of ecosystem functions within one historic native and two non-native species assemblages under different grazing intensities to address how different species assemblages vary in provisioning, regulatory and supporting ecosystem services. Forage production was reduced in one non-native assemblage (medusahead). Cultural ecosystem services, such as native species diversity, were inherently lower in both non-native assemblages, whereas most other services were maintained across grazing intensities. All systems provided similar ecosystem services under the highest grazing intensity treatment, which simulated unsustainable grazing intensity. We suggest that applying a more comprehensive ecosystem framework that considers multiple ecosystem services to evaluate new emerging ecosystems is a valuable tool to determine management goals and how to intervene in a changing ecosystem.     

Species effects on resource supply rates: do they influence competitive interactions?

We examined two ways in which species effects on nitrogen supply rates could influence species competitive interactions and lead to coexistence between species with very different patterns of resource use. First, through differential effects on the rates of resource cycling, species may modify resources in divergent directions to better tolerate neighbors under self-modified conditions (tolerance modification). Second, plant-induced shifts in resource supply could enhance the degree to which species can suppress individuals (suppression modification). We addressed these hypothesized mechanisms in the non-successional moist meadow alpine tundra, which is codominated by Acomastylis rossii, a slow-growing roseaceous forb with high nutrient retention that is associated with slow rates of nitrogen supply, and Deschampsia caespitosa, a graminoid with rapid growth potential that is associated with fast rates of nitrogen (N) supply. Neighbors inhibited Acomastylis transplants (the species with high N retention) less than Deschampsia transplants (the species with rapid potential growth) in both neighborhood types, and hence neighborhood type did not influence species tolerance to neighbors. Likewise, Acomastylis neighborhoods inhibited transplant growth more than Deschampsia neighborhoods regardless of transplant species identity. When N supply rates were enhanced through repeated N additions in the two neighborhood types or in the presence of each species’ litter, the inhibitory effects associated with Acomastylis were offset to a greater degree than those associated with Deschampsia, as predicted by the suppression modification. These effects appeared to be density or size-dependent, with few effects observed at low density. Our results suggest that species effects on supply rates may influence competitive interactions, particularly if these effects are complemented by other sources of temporal or spatial variation such as pulses in resource availability. This revised version was published online in June 2006 with corrections to the Cover Date.     

Neotropical rainforest restoration: comparing passive,plantation and nucleation approaches

Neotropical rainforests are global biodiversity hotspots and are challenging to restore. A core part of this challenge is the very long recovery trajectory of the system: recovery of structure can take 20–190 years, species composition 60–500 years, and reestablishment of rare/endemic species thousands of years. Passive recovery may be fraught with instances of arrested succession, disclimax or emergence of novel ecosystems. In these cases, active restoration methods are essential to speed recovery and set a desired restoration trajectory. Tree plantation is the most common active approach to reestablish a high density of native tree species and facilitate understory regeneration. While this approach may speed the successional trajectory, it may not achieve, and possibly inhibit, a long-term restoration trajectory towards the high species diversity characteristic of these forests. A range of nucleation techniques (e.g., tree island planting) are important restoration options: although they may not speed recovery of structure as quickly as plantations, their emphasis on natural regeneration processes may enable greater and more natural patterns of diversity to develop. While more work needs to be done to compare forest restoration techniques in different environmental contexts, it appears that nucleation and, at times, passive restoration may best preserve the diverse legacy of these forested systems (both with lower costs). An integrated approach using both plantation productivity but also the natural functions associated with nucleation may develop composition and diversity trajectory desired in Neotropical conservation efforts.     

Strong feeding preference of an exotic generalist herbivore for an exotic forb: a case of invasional antagonism

Many hypotheses dealing with the success of invasive plant species concern plant–herbivore interactions. The invasional meltdown and enemy inversion hypotheses suggest that non-native herbivores may indirectly facilitate the invasion of a non-native plant species by either favorably changing environmental conditions or reducing competition from native plant species. Our objective was to determine the role of herbivory by the non-native snail Otala lactea in structuring California grassland communities. We conducted two experiments to examine the feeding preferences of O. lactea for eight representative grassland species. Overall, O. lactea preferred Brassica nigra, a non-native forb, over all other species tested. Field monocultures of B. nigra supported significantly higher snail densities than monocultures of any of the other species tested. O. lactea also preferred B. nigra over all other species tested in controlled laboratory feeding trials. However, based on trait comparisons of each of the eight grassland species, we cannot pinpoint the preference for B. nigra to a basic nutritional requirement on the part of the herbivore or an allocation to defense on the part of the plants. Our study provides evidence for an antagonistic relationship between a non-native herbivore and a non-native plant species in their invasive range. We term this relationship “invasional antagonism”.     

The effects of gap creation on competitive interactions: separating changes in overall intensity from relative rankings

A central assumption of disturbance ecology equates gap creation with the reduction of overall competitive intensity. I develop the idea that gap creation, due to changes in addition to biomass removal, could also alter the relative competitive rankings among species. I present a quantitative review using meta-analysis in which I separate the effect of gap creation from the effect of competition for species characteristic of different disturbance regimes. Twenty-one studies (with a total of 136 comparisons) directly examined species competitive abilities under gap (disturbed) and matrix (undisturbed) conditions. Overall competitive intensity generally declined in gap conditions, although species characteristic of gap areas responded more strongly to gap creation than did species characteristic of undisturbed matrix areas. Under matrix conditions, matrix species were less affected by competition than gap species, supporting one widespread assumption. However, under gap conditions, matrix and gap species were inhibited to a similar degree by neighbor biomass. These results suggest that gap creation, in addition to decreasing overall competitive intensity, may affect species competitive rankings, possibly due to changes in the environment where the interactions occur.     

Frequency-dependence stabilizes competitive interactions among four annual plants

It is the combination of large fitness differences and strong stabilizing mechanisms that often constitute niche-based explanations for species abundance patterns. Despite the importance of this assumption to much of community ecology, empirical evidence is surprisingly limited. Empirical tests are critical because many abundance patterns are also consistent with neutral-based alternatives (that assume no fitness differences or stabilization). We quantified interactions of four annual grassland species in two-species mixtures at varying frequencies. We found evidence of strong negative frequency-dependent stabilization, where scaled population growth rates increased with decreasing frequency for all four species. There was also a consistent competitive hierarchy among these species indicative of strong fitness differences that, in most cases, suggested potential competitive exclusion despite the observed strong stabilization.     

Environmental gradients determine the potential for ecosystem engineering effects

Understanding processes that determine biodiversity is a fundamental challenge in ecology. At the landscape scale, physical alteration of ecosystems by organisms, called ecosystem engineering, enhances biodiversity worldwide by increasing heterogeneity in resource conditions and enhancing species coexistence across engineered and non‐engineered habitats. Engineering–diversity relationships can vary along environmental gradients due to changes in the amount of physical structuring created by ecosystem engineering, but it is unclear how this variation is influenced by the responsiveness of non‐structural abiotic properties to engineering. Here we show that environmental gradients determine the capacity for engineering to alter resource availability and species diversity, independent of the magnitude of structural change produced by engineering. We created an experimental rainfall gradient in an arid grassland where rodents restructure soils by constructing large, long‐lasting burrows. We found that greater rainfall increased water availability and productivity in both burrow and inter‐burrow habitats, causing a decline in local (alpha) plant diversity within both of these habitats. However, increased rainfall also resulted in greater differences in soil resources between burrow and inter‐burrow habitats, which increased species turnover (beta diversity) across habitats and stabilized landscape‐level (gamma) diversity. These responses occurred regardless of rodent presence and without changes in the extent of physical alteration of soils by rodents. Our results suggest that environmental gradients can influence the effects of ecosystem engineering in maintaining biodiversity via resource heterogeneity and species turnover. In an era of rapid environmental change, accounting for this interaction may be critical to conservation and management.     

Vegetation change at high elevation: scale dependence and interactive effects on Niwot Ridge

Background: High-elevation mountain systems may be particularly responsive to climate change.

Aims: Here we investigate how changes along elevation gradients in mountain systems can aid in predicting vegetation distributional changes in time, focusing on how changing climatic controls affect meso-scale transitions at the lower and upper boundaries of alpine vegetation (with forest and subnival zones, respectively) as well as micro-scale transitions among plant communities within the alpine belt. We focus on climate-related drivers, particularly in relation to climate change, but also consider how species interactions, dispersal and responses to disturbance may influence plant responses to these abiotic drivers.

Results: Empirical observations and experimental studies indicate that changing climatic controls influence both meso-scale transitions at the upper and lower boundaries of alpine vegetation and micro-scale transitions among plant communities within tundra. Micro-scale heterogeneity appears to buffer response in many cases, while interactions between climate and other changes may often accelerate change.

Conclusions: Interactions with microtopography and larger edaphic gradients have the capacity to both facilitate rapid changes and reinforce stability, and that these interactions will affect the responsiveness of vegetation to climate change at different spatial scales.     

Neue diterpene aus der subtribus espeletiinae

The investigation of several species from the newly created subtribe Espeletiinae afforded in addition to known compounds seven new diterpenes and two norditerpenes. The structures were elucidated by intensive ¹H-NMR studies. The chemotaxonomic situation is discussed briefly.