Erik T. Aschehoug

Positive interactions among alpine plants increase with stress

Plants can have positive effects on each other. For example, the accumulation of nutrients, provision of shade, amelioration of disturbance, or protection from herbivores by some species can enhance the performance of neighbouring species. Thus the notion that the distributions and abundances of plant species are independent of other species may be inadequate as a theoretical underpinning for understanding species coexistence and diversity. But there have been no large-scale experiments designed to examine the generality of positive interactions in plant communities and their importance relative to competition. Here we show that the biomass, growth and reproduction of alpine plant species are higher when other plants are nearby. In an experiment conducted in subalpine and alpine plant communities with 115 species in 11 different mountain ranges, we find that competition generally, but not exclusively, dominates interactions at lower elevations where conditions are less physically stressful. In contrast, at high elevations where abiotic stress is high the interactions among plants are predominantly positive. Furthermore, across all high and low sites positive interactions are more important at sites with low temperatures in the early summer, but competition prevails at warmer sites.

Plant behavioural ecology: dynamic plasticity in secondary metabolites

Behaviour is in part the ability to respond rapidly and reversibly in response to environmental stimuli during the lifetime of an individual. Plants and animals both exhibit behaviour, but plant behaviour is most often examined in the context of morphologically plastic growth. Rapid and reversible secondary metabolite production and release is also a key mechanism by which plants behave. Here, we review plant biochemical plasticity as plant behaviour, and explicitly focus on evidence for responses that display rapid induction, reversibility and ecological relevance. Rapid induction and attenuation of plant secondary metabolites occur as chemically mediated root foraging, plant defence, allelochemistry and to regulate mutualistic relationships. We describe a wealth of information on the induction of various plant biochemical responses to environmental stimuli but found a limited body of literature on the reversibility of induced biochemical responses. Understanding the full cycle of dynamic plasticity in secondary metabolites is an important niche for future research. Biochemical behaviours extend beyond the plant kingdom; however, they clearly illustrate the capacity for plants to behave in ways that closely mirror the classic definitions and research approaches applied to behaviour in animals.

A framework to assess evolutionary responses to anthropogenic light and sound

Human activities have caused a near-ubiquitous and evolutionarily-unprecedented increase in environmental sound levels and artificial night lighting. These stimuli reorganize communities by interfering with species-specific perception of time-cues, habitat features, and auditory and visual signals. Rapid evolutionary changes could occur in response to light and noise, given their magnitude, geographical extent, and degree to which they represent unprecedented environmental conditions. We present a framework for investigating anthropogenic light and noise as agents of selection, and as drivers of other evolutionary processes, to influence a range of behavioral and physiological traits such as phenological characters and sensory and signaling systems. In this context, opportunities abound for understanding contemporary and rapid evolution in response to human-caused environmental change.

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebes competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.

Fungal endophyte increases the allelopathic effects of an invasive forb.

Abstract Endophytic plant symbionts can have powerful effects on the way their hosts interact with pathogens, competitors, and consumers. The presence of endophytes in plants can alter food webs, community composition and ecosystem processes, suggesting that endophyte-plant symbioses may represent unique forms of extended phenotypes. We tested the impact of the fungal endophyte Alternaria alternata (phylotype CID 120) on the allelopathic effect of the invasive forb Centaurea stoebe when in competition with the North American native bunchgrass Koeleria macrantha in a greenhouse competition experiment. The allelopathic effect of C. stoebe on K. macrantha when infected with the fungal endophyte was more than twice that of endophyte-free C. stoebe. However, this allelopathic effect was a small part of the very large competitive effect of C. stoebe on K. macrantha in all treatments, likely because of the priority effects in our experimental design. To our knowledge, these results are the first experimental evidence for a symbiotic relationship between plants and fungal endophytes affecting allelopathic interactions between competing plants, and thus provide insight into the mechanisms by which fungal endophytes may increase the competitive ability of their hosts.

Diversity Increases Indirect Interactions, Attenuates the Intensity of Competition, and Promotes Coexistence

A fundamental assumption of coexistence theory is that competition inevitably decreases species diversity. Consequently, in the quest to understand the ecological regulators of diversity, there has been a great deal of focus on processes with the potential to reduce competitive exclusion. However, the notion that competition must decrease diversity is largely based on the outcome of two-species interaction experiments and models, despite the fact that species rarely interact only in pairs in natural systems. In a field experiment, we found that competition among native perennial plants in multispecies assemblages was far weaker than competition between those same species in pairwise arrangements and that indirect interactions appeared to weaken direct competitive effects. These results suggest that community assembly theory based on pairwise approaches may overestimate the strength of competition and likelihood of competitive exclusion in species-rich communities. We also found that Centaurea stoebe, a North American invader, retained strong competitive effects when competing against North American natives in both pairwise and multispecies assemblages. Our experimental results support an emerging body of theory suggesting that complex networks of competing species may generate strong indirect interactions that can maintain diversity and that ecological differentiation may not be necessary to attenuate competition.

Morphological variability in tree root architecture indirectly affects coexistence among competitors in the understory.

Interactions between plants can have strong effects on community structure and function. Variability in the morphological, developmental, physiological, and biochemical traits of plants can influence the outcome of plant interactions and thus have important ecological consequences. However, the ecological ramifications of trait variability in plants are poorly understood and have rarely been tested in the field. We experimentally tested the effects of morphological variation in root architecture of Quercus douglasii trees in the field on interactions between understory plants and community composition. Our results indicate that variability among Q. douglasii tree root systems initiates a striking reversal in the competitive effects of dominant understory grass species on a less common species. Trees with a deep-rooted morphology facilitated exotic annual grasses and these annual grasses, in turn, competitively excluded the native perennial bunchgrass, Stipapulchra. In contrast, Q. douglasii trees with shallow-rooted morphologies directly suppressed the growth of exotic annual grasses and indirectly released S. pulchra individuals from competition with these annual grasses. Morphological variation in the root architecture of Q. douglasii created substantial conditionality in the outcomes of competition among species which enhanced the potential for indirect interactions to sustain coexistence and increase community diversity.

Models of Experimental Competitive Intensities Predict Home and Away Differences in Invasive Impact and the Effects of an Endophytic Mutualist

Understanding the role of competition in the organization of communities is limited in part by the difficulty of extrapolating the outcomes of small-scale experiments to how such outcomes might affect the distribution and abundance of species. We modeled the community-level outcomes of competition, using experimentally derived competitive effects and responses between an exotic invasive plant, Centaurea stoebe, and species from both its native and nonnative ranges and using changes in these effects and responses elicited by experimentally establishing symbioses between C. stoebe and fungal endophytes. Using relative interaction intensities (RIIs) and holding other life-history factors constant, individual-based and spatially explicit models predicted competitive exclusion of all but one North American species but none of the European species, regardless of the endophyte status of C. stoebe. Concomitantly, C. stoebe was eliminated from the models with European natives but was codominant in models with North American natives. Endophyte symbiosis predicted increased dominance of C. stoebe in North American communities but not in European communities. However, when experimental variation was included, some of the model outcomes changed slightly. Our results are consistent with the idea that the effects of competitive intensity and mutualisms measured at small scales have the potential to play important roles in determining the larger-scale outcomes of invasion and that the stabilizing indirect effects of competition may promote species coexistence.

Habitat restoration affects immature stages of a wetland butterfly through indirect effects on predation

Habitat loss worldwide has led to the widespread use of restoration practices for the recovery of imperiled species. However, recovery success may be hampered by focusing on plant communities, rather than the complex suite of direct and indirect interactions among trophic levels that occur in natural systems. Through a factorial field experiment, we tested the effects of wetland restoration on egg and juvenile survival of a locally rare butterfly, Satyrodes appalachia, via tree removal and damming. Tree removal more than tripled S. appalachia host plant abundance, but neither restoration action directly affected S. appalachia egg and juvenile survival. Instead, we found strong indirect effects of habitat manipulation on S. appalachia egg and juvenile survival that were mediated through predation. The interaction of tree removal and damming significantly decreased predation of S. appalachia eggs relative to each treatment alone. Damming alone had a significant positive indirect effect on the survival of S. appalachia juveniles, likely because increases in standing water reduced predator access. Our results emphasize the need for experiments that evaluate the demographic responses of imperiled species to habitat restoration prior to management action and quantify potential indirect effects mediated through higher trophic levels.

Habitat Restoration as a Recovery Tool for a Disturbance-Dependent Butterfly, The Endangered St. Francis’ Satyr

Habitat restoration is a key management strategy for species that experience loss, fragmentation or degradation of their habitat. The St. Francis’ satyr is a federally endangered butterfly found only on Ft. Bragg Army Installation in North Carolina that depends on frequent disturbance to maintain its preferred habitat, ephemeral wetlands. Over the course of 4 years, we restored and maintained critical habitat for the St. Francis’ satyr through a combination of hardwood removal and inundation via artificial dams to mimic natural beaver and fire disturbance. Here we present our insights into the challenges of creating and maintaining restored habitat for a species dependent on frequent disturbance, the results of experimental testing of demographic success, and the complexities in carrying out a captive rearing program. We discuss how species that are dependent on disturbance created habitat pose a unique challenge in creating and implementing a long-term plan to maintain a dynamic landscape.