No abstract

When introduced to new habitats by humans, some plant species become much more dominant. This is primarily attributed to escape from specialist consumers. Release from these specialist enemies is also thought by some to lead to the evolution of increased competitive ability, driven by a decrease in the plant's resource allocation to consumer defense and an increase in allocation to size or fecundity. Here, we discuss a new theory for invasive success – the “novel weapons hypothesis”. We propose that some invaders transform because they possess novel biochemical weapons that function as unusually powerful allelopathic agents, or as mediators of new plant–soil microbial interactions. Root exudates that are relatively ineffective against their natural neighbors because of adaptation, may be highly inhibitory to newly encountered plants in invaded communities. In other words, the novel weapons of some plant invaders provide them with an advantage that may arise from differences in the regional coevolutionary trajectories of plant communities. Furthermore, the selective advantage of possessing a novel weapon may result in rapid evolution of that weapon – for example, the production of greater quantities of allelopathic or antimicrobial root exudates. Direct selection of competitive traits provides an alternative to the “grow versus defend” trade‐offs that underpin the theory of the evolution of increased competitive ability.

Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a 'safety net' sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

The relative importance of allelopathy and resource competition in plant-plant interactions has been vigorously debated but seldom tested. We used activated carbon to manipulate the effects of root exudates of Centaurea maculosa, a noxious weed in much of western North America, on root elongation rates and growth of the native bunchgrass Festuca idahoensis in order to investigate the relative importance of allelopathy in the total interference of Centaurea. In root observation chambers, Festuca root elongation rates decreased to ≈50% of the control, beginning 4 days before contacting Centaurea roots in silica sand. However, when activated carbon, which has a high affinity for adsorbing to organic compounds, was added to the sand the effects of Centaurea roots on Festuca root elongation were reduced. In other experiments, Festuca plants were 50% smaller when grown with Centaurea than with conspecifics in pure silica sand. However, Festuca grown with Centaurea in mixtures of sand and activated carbon were 85% larger than Festuca grown with Centaurea in silica sand without carbon. These results suggest that allelopathy accounts for a substantial proportion of the total interference of Centaurea on Festuca, shifting the balance of competition in favor of Centaurea. However, Centaurea outperformed Festuca even in the presence of activated carbon, demonstrating the importance of the combined roles of resource competition and allelopathy.

Summary1 Exotic plant invasions often cause high mortality in native populations and therefore have the potential to be a powerful selective force. 2 We found that surviving individuals from North American communities that have experienced extensive invasion by Centaurea maculosa have higher tolerances to the Eurasian invader than individuals from communities that did not experience invasion. 3 Some native species grown from the seed of individuals that survived Centaurea invasion were more resistant to the general competitive effects of Centaurea , the root exudates from Centaurea , and to a chemical specific to the root exudates of Centaurea ( ± )-catechin. 4 Although these results may be confounded by maternal effects, they provide initial evidence that native plant species may evolve to tolerate the effects of an exotic invader, and in particular an invader's novel allelochemistry. 5 Such effects may have long-term implications for plant invasions and the organization of plant communities. Evolved tolerance may ultimately contribute to coexistence among natives and invaders, with our results suggesting that natural plant communities may be more coevolved and less individualistic than currently thought.

The natural enemies hypothesis has led to a number of ideas by which invaders might evolve superior competitive ability. In this context, we compared growth, reproduction, competitive effect, competitive response, and defense capabilities between invasive North American populations of Centaurea maculosa and populations in Europe, where the species is native. We found that Centaurea from North America were larger than plants from European populations. North American Centaurea also demonstrated stronger competitive effects and responses than European Centaurea. However, competitive superiority did not come at a cost to herbivore defense. North American plants were much better defended against generalist insect herbivores and slightly better defended against specialists. North Americans showed a stronger inhibitory effect on the consumers (resistance) and a better ability to regrow after attack by herbivores (tolerance). Better defense by North Americans corresponded with higher constitutive levels of a biochemical defense compound precursor, tougher leaves, and more leaf trichomes than Europeans. North American F1 progeny of field collected lines retained the traits of larger size and greater leaf toughness suggesting that genetic differences, rather than maternal effects, may be the cause of intercontinental differences, but these sample sizes were small. Our results suggest that the evolution of increased competitive ability may not always be driven by physiological trade‐offs between the allocation of energy or resources to growth or to defense. Instead, we hypothesize that Centaurea maculosa experiences strong directional selection on novel competitive and defense traits in its new range.

Biotic interactions can shape phylogenetic community structure (PCS). However, we do not know how the asymmetric effects of foundation species on communities extend to effects on PCS. We assessed PCS of alpine plant communities around the world, both within cushion plant foundation species and adjacent open ground, and compared the effects of foundation species and climate on alpha (within-microsite), beta (between open and cushion) and gamma (open and cushion combined) PCS. In the open, alpha PCS shifted from highly related to distantly related with increasing potential productivity. However, we found no relationship between gamma PCS and climate, due to divergence in phylogenetic composition between cushion and open sub-communities in severe environments, as demonstrated by increasing phylo-beta diversity. Thus, foundation species functioned as micro-refugia by facilitating less stress-tolerant lineages in severe environments, erasing a global productivity - phylogenetic diversity relationship that would go undetected without accounting for this important biotic interaction.

Summary1 Plant allelopathic compounds may have other roles than interspecific interference. We investigated whether (±)-catechin, a phytotoxin exuded from the roots of the exotic invader Centaurea maculosa (spotted knapweed), is also one of the factors that regulates C. maculosa recruitment. 2 Adding activated carbon, which adsorbs organic compounds, to soil around C. maculosa adults in the field increased seedling density by 78% 25 days after sowing, and by 34% 32 days after sowing, suggesting that soil-borne compounds inhibited or delayed recruitment. 3 Analysis of field soils near mature C. maculosa revealed that they can contain exceptionally high (±)-catechin concentrations (mean = 1.55 mg g −1 dry soil, with 60% of samples containing ≥ 1.0 mg (±)-catechin g −1 ). 4 In laboratory experiments, treatment with ≥ 1.0 mg (±)-catechin mL −1 reduced seedling root elongation by > 50%, indicating that field concentrations are sufficient to inhibit C. maculosa recruitment. Provided that 10% methanol was used to maintain (±)-catechin in solution for > 1 day, treatment with ≥ 1.0 mg mL −1 also reduced C. maculosa germination by > 70%. 5 (±)-Catechin maintained in solution with methanol did not significantly reduce C. maculosa seed survival, suggesting that inhibition of germination was due, at least in part, to delayed germination rather than to seed mortality. 6 Depending on the solubility of (±)-catechin in soil and on the duration of its effects on recruitment, C. maculosa may avoid intraspecific competition or regulate the timing of seedling establishment by reducing seedling growth or postponing germination in response to its own phytotoxin. 7 Chemical regulation of C. maculosa recruitment, as demonstrated here, suggests a dual role of (±)-catechin as an allelochemical and an autoinhibitor. The potential for a single plant root exudate to influence both inter-and intraspecific interactions emphasizes the complex effects that plant secondary metabolites may have on plant population and community structure.