Phenotypic and genetic differentiation between native and introduced plant populations

Plant invasions often involve rapid evolutionary change. Founder effects, hybridization, and adaptation to novel environments cause genetic differentiation between native and introduced populations and may contribute to the success of invaders. An influential idea in this context has been the Evolution of Increased Competitive Ability (EICA) hypothesis. It proposes that after enemy release plants rapidly evolve to be less defended but more competitive, thereby increasing plant vigour in introduced populations. To detect evolutionary change in invaders, comparative studies of native versus introduced populations are needed. Here, we review the current empirical evidence from: (1) comparisons of phenotypic variation in natural populations; (2) comparisons of molecular variation with neutral genetic markers; (3) comparisons of quantitative genetic variation in a common environment; and (4) comparisons of phenotypic plasticity across different environments. Field data suggest that increased vigour and reduced herbivory are common in introduced plant populations. In molecular studies, the genetic diversity of introduced populations was not consistently different from that of native populations. Multiple introductions of invasive plants appear to be the rule rather than the exception. In tests of the EICA hypothesis in a common environment, several found increased growth or decreased resistance in introduced populations. However, few provided a full test of the EICA hypothesis by addressing growth and defence in the same species. Overall, there is reasonable empirical evidence to suggest that genetic differentiation through rapid evolutionary change is important in plant invasions. We discuss conceptual and methodological issues associated with cross-continental comparisons and make recommendations for future research. When testing for EICA, greater emphasis should be put on competitive ability and plant tolerance. Moreover, it is important to address evolutionary change in characteristics other than defence and growth that could play a role in plant invasion

Mechanisms of Chinese tallow (Triadica sebifera) invasion and their management implications – A review

Ecosystems are under increasing stress from environmental change, including invasion by non-native species that can disrupt ecological processes and functions. Chinese tallow [Triadica sebifera (L.) Small] is a highly invasive tree species in southeastern US forests, prairies, and wetlands, and effectively managing this invasive species is a significant challenge for scientists and land managers. In this review, we synthesize the literature on invasion ecology and management of Chinese tallow. Our review suggests that the invaded range of Chinese tallow is currently limited by dispersal in many areas and by low temperatures and low soil moisture, and by high soil salinity and frequent flooding in others, but these barriers may be overcome by increased dispersal, phenotypic plasticity, and/or rapid evolution. Invasions by Chinese tallow are facilitated by both the invasiveness of the species and the invasibility of the recipient communities. Invasiveness of Chinese tallow has been attributed to fast growth, high fecundity, a persistent seed bank, aggressive resprouting, abiotic stress tolerance, and the ability to transform fire maintained ecosystems. Some of these traits may be enhanced in invasive populations. Anthropogenic and natural disturbances, lack of herbivore pressure, and facilitation by soil microbes enhance the intensity of Chinese tallow invasions. Biological control of Chinese tallow is being developed. Treatments such as herbicides, prescribed fire, and mechanical control can effectively control Chinese tallow at the local scale. A combination of these treatments improves results. However, a proactive management approach would simultaneously achieve invasion control and promote subsequent ecological restoration, especially in the context of legacy effects, secondary invasions, and/or variable ecosystem responses to different control treatments. Future research should clarify the roles of species invasiveness and community invasibility, increase our understanding of the effects of Chinese tallow in invaded communities, and develop viable management regimes that are effective in both controlling or reducing the probability of Chinese tallow invasion and restoring desired native communities

Species‐specific plant‐mediated effects between herbivores converge at high damage intensity

Plants are often exposed to multiple herbivores and densities of these attackers (or corresponding damage intensities) often fluctuate greatly in the field. Plant-mediated interactions vary among herbivore species and with changing feeding intensity, but little is known about how herbivore identity and density interact to determine plant responses and herbivore fitness. Here, we investigated this question using Triadica sebifera (tallow) and two common and abundant specialist insect herbivores, Bikasha collaris (flea beetle) and Heterapoderopsis bicallosicollis (weevil). By manipulating densities of leaf-feeding adults of these two herbivore species, we tested how variations in the intensity of leaf damage caused by flea beetle or weevil adults affected the performance of root-feeding flea beetle larvae and evaluated the potential of induced tallow root traits to predict flea beetle larval performance. We found that weevil adults consistently decreased the survival of flea beetle larvae with increasing leaf damage intensities. In contrast, conspecific flea beetle adults increased their larval survival at low damage then decreased larval survival at high damage, resulting in a unimodal pattern. Chemical analyses showed that increasing leaf damage from weevil adults linearly decreased root carbohydrates and increased root tannin, whereas flea beetle adults had opposite effects as weevil adults at low damage and similar effects as them at high damage. Furthermore, across all feeding treatments, flea beetle larval survival correlated positively with concentrations of carbohydrates and negatively with concentration of tannin, suggesting that root primary and secondary metabolism might underlie the observed effects on flea beetle larvae. Our study demonstrates that herbivore identity and density interact to determine systemic plant responses and plant-mediated effects on herbivores. In particular, effects are species-specific at low densities, but converge at high densities. These findings emphasize the importance of considering herbivore identity and density simultaneously when investigating factors driving plant-mediated interactions between herbivores, which advances our understanding of the structure and composition of herbivore communities and terrestrial food webs

Non-Native Plant Litter Enhances Soil Carbon Dioxide Emissions in an Invaded Annual Grassland

Litter decomposition is a fundamental ecosystem process in which breakdown and decay of plant detritus releases carbon and nutrients. Invasive exotic plants may produce litter that differs from native plant litter in quality and quantity. Such differences may impact litter decomposition and soil respiration in ways that depend on whether exotic and native plant litters decompose in mixtures. However, few field experiments have examined how exotic plants affect soil respiration via litter decomposition. Here, we conducted an in situ study of litter decomposition of an annual native grass (Eragrostis pilosa), a perennial exotic forb (Alternanthera philoxeroides), and their mixtures in an annual grassland in China to examine potential invasion effects on soil respiration. Alternanthera litter decomposed faster than Eragrostis litter when each was incubated separately. Mass loss in litter mixes was more rapid than predicted from rates in single species bags (only 35% of predicted mass remained at 8 months) showing synergistic effects. Notably, exotic plant litter decomposition rate was unchanged but native plant litter decomposition rate was accelerated in mixtures (decay constant k = 0.20 month(−1)) compared to in isolation (k = 0.10 month(−1)). On average, every litter type increased soil respiration compared to bare soil from which litter was removed. However, the increases were larger for mixed litter (1.82 times) than for Alternanthera litter (1.58 times) or Eragrostis litter (1.30 times). Carbon released as CO(2) relative to litter carbon input was also higher for mixed litter (3.34) than for Alternathera litter (2.29) or Eragrostis litter (1.19). Our results indicated that exotic Alternanthera produces rapidly decomposing litter which also accelerates the decomposition of native plant litter in litter mixtures and enhances soil respiration rates. Thus, this exotic invasive plant species will likely accelerate carbon cycling and increase soil respiration even at intermediate stages of invasion in these annual grasslands

Timing of Favorable Conditions, Competition and Fertility Interact to Govern Recruitment of Invasive Chinese Tallow Tree in Stressful Environments

The rate of new exotic recruitment following removal of adult invaders (reinvasion pressure) influences restoration outcomes and costs but is highly variable and poorly understood. We hypothesize that broad variation in average reinvasion pressure of Triadica sebifera (Chinese tallow tree, a major invader) arises from differences among habitats in spatiotemporal availability of realized recruitment windows. These windows are periods of variable duration long enough to permit establishment given local environmental conditions. We tested this hypothesis via a greenhouse mesocosm experiment that quantified how the duration of favorable moisture conditions prior to flood or drought stress (window duration), competition and nutrient availability influenced Triadica success in high stress environments. Window duration influenced pre-stress seedling abundance and size, growth during stress and final abundance; it interacted with other factors to affect final biomass and germination during stress. Stress type and competition impacted final size and biomass, plus germination, mortality and changes in size during stress. Final abundance also depended on competition and the interaction of window duration, stress type and competition. Fertilization interacted with competition and stress to influence biomass and changes in height, respectively, but did not affect Triadica abundance. Overall, longer window durations promoted Triadica establishment, competition and drought (relative to flood) suppressed establishment, and fertilization had weak effects. Interactions among factors frequently produced different effects in specific contexts. Results support our ‘outgrow the stress’ hypothesis and show that temporal availability of abiotic windows and factors that influence growth rates govern Triadica recruitment in stressful environments. These findings suggest that native seed addition can effectively suppress superior competitors in stressful environments. We also describe environmental scenarios where specific management methods may be more or less effective. Our results enable better niche-based estimates of local reinvasion pressure, which can improve restoration efficacy and efficiency by informing site selection and optimal Management

Bacterial and fungal inhibitor interacted impacting growth of invasive Triadica sebifera and soil N2O emissions

Plant invasions affect biodiversity and seriously endanger the stability of ecosystems. Invasive plants show strong adaptability and growth advantages but are influenced by various factors. Soil bacteria and fungi are critical to plant growth and are important factors affecting plant invasions. Plant invasions also affect soil N2O emissions, but the effects of invasive plants from different population origins on N2O emissions and their microbial mechanisms are not clear. In this experiment, we grew Triadica sebifera from native (China) and invasive (USA) populations with or without bacterial (streptomycin) and/or fungal (iprodione) inhibitors in a factorial experiment in which we measured plant growth and soil N2O emissions of T. sebifera. Plants from invasive populations had higher leaf masses than those from native populations when soil bacteria were not inhibited (with or without fungal inhibition) which might reflect that they are more dependent on soil bacteria. Cumulative N2O emissions were higher for soils with invasive T. sebifera than those with a plant from a native population. Bacterial inhibitor application reduced cumulative N2O emissions but reductions were larger with application of the fungal inhibitor either alone or in combination with the bacterial inhibitor. This suggests that fungi play a strong role in plant performance and soil N2O emissions. Therefore, it is important to further understand the effects of soil microorganisms on the growth of T. sebifera and soil N2O emissions to provide a more comprehensive scientific basis for understanding the causes and consequences of plant invasions

Evolutionary dynamics of tree invasions: complementing the unified framework for biological invasions

Evolutionary processes greatly impact the outcomes of biological invasions. An extensive body of research suggests that invasive populations often undergo phenotypic and ecological divergence from their native sources. Evolution also operates at different and distinct stages during the invasion process. Thus, it is important to incorporate evolutionary change into frameworks of biological invasions because it allows us to conceptualize how these processes may facilitate or hinder invasion success. Here, we review such processes, with an emphasis on tree invasions, and place them in the context of the unified framework for biological invasions. The processes and mechanisms described are pre-introduction evolutionary history, sampling effect, founder effect, genotype-by-environment interactions, admixture, hybridization, polyploidization, rapid evolution, epigenetics and second-genomes. For the last, we propose that co-evolved symbionts, both beneficial and harmful, which are closely physiologically associated with invasive species, contain critical genetic traits that affect the evolutionary dynamics of biological invasions. By understanding the mechanisms underlying invasion success, researchers will be better equipped to predict, understand and manage biological invasions

The Effect of Diet and Opponent Size on Aggressive Interactions Involving Caribbean Crazy Ants (Nylanderia fulva)

Biotic interactions are often important in the establishment and spread of invasive species. In particular, competition between introduced and native species can strongly influence the distribution and spread of exotic species and in some cases competition among introduced species can be important. The Caribbean crazy ant, Nylanderia fulva, was recently introduced to the Gulf Coast of Texas, and appears to be spreading inland. It has been hypothesized that competition with the red imported fire ant, Solenopsis invicta, may be an important factor in the spread of crazy ants. We investigated the potential of interspecific competition among these two introduced ants by measuring interspecific aggression between Caribbean crazy ant workers and workers of Solenopsis invicta. Specifically, we examined the effect of body size and diet on individual-level aggressive interactions among crazy ant workers and fire ants. We found that differences in diet did not alter interactions between crazy ant workers from different nests, but carbohydrate level did play an important role in antagonistic interactions with fire ants: crazy ants on low sugar diets were more aggressive and less likely to be killed in aggressive encounters with fire ants. We found that large fire ants engaged in fewer fights with crazy ants than small fire ants, but fire ant size affected neither fire ant nor crazy ant mortality. Overall, crazy ants experienced higher mortality than fire ants after aggressive encounters. Our findings suggest that fire ant workers might outcompete crazy ant workers on an individual level, providing some biotic resistance to crazy ant range expansion. However, this resistance may be overcome by crazy ants that have a restricted sugar intake, which may occur when crazy ants are excluded from resources by fire ants

Facilitation and Competition among Invasive Plants: A Field Experiment with Alligatorweed and Water Hyacinth

Ecosystems that are heavily invaded by an exotic species often contain abundant populations of other invasive species. This may reflect shared responses to a common factor, but may also reflect positive interactions among these exotic species. Armand Bayou (Pasadena, TX) is one such ecosystem where multiple species of invasive aquatic plants are common. We used this system to investigate whether presence of one exotic species made subsequent invasions by other exotic species more likely, less likely, or if it had no effect. We performed an experiment in which we selectively removed exotic rooted and/or floating aquatic plant species and tracked subsequent colonization and growth of native and invasive species. This allowed us to quantify how presence or absence of one plant functional group influenced the likelihood of successful invasion by members of the other functional group. We found that presence of alligatorweed (rooted plant) decreased establishment of new water hyacinth (free-floating plant) patches but increased growth of hyacinth in established patches, with an overall net positive effect on success of water hyacinth. Water hyacinth presence had no effect on establishment of alligatorweed but decreased growth of existing alligatorweed patches, with an overall net negative effect on success of alligatorweed. Moreover, observational data showed positive correlations between hyacinth and alligatorweed with hyacinth, on average, more abundant. The negative effect of hyacinth on alligatorweed growth implies competition, not strong mutual facilitation (invasional meltdown), is occurring in this system. Removal of hyacinth may increase alligatorweed invasion through release from competition. However, removal of alligatorweed may have more complex effects on hyacinth patch dynamics because there were strong opposing effects on establishment versus growth. The mix of positive and negative interactions between floating and rooted aquatic plants may influence local population dynamics of each group and thus overall invasion pressure in this watershed