Bernd Blossey

Before, During and After: The Need for Long-term Monitoring in Invasive Plant Species Management

The invasion of non-indigenous plants is considered one of the primary threats to rare and endangered species as well as to the integrity and function of North American ecosystems. However, many of the suspected negative ecosystem impacts are based on anecdotal evidence. For example, there is almost unanimous agreement among natural resource managers of the detrimental ecological impacts of species such as Lythrum salicaria (purple loosestrife), Phragmites australis (common reed) and Alliaria petiolata (garlic mustard) but convincing documentation is scarce. Experimental and theoretical ecology predicts large ecosystem impacts of the most widespread invasive species. However, it is difficult to prioritize control of species that occur at intermediate densities. Long-term monitoring before and during the invasion as well as before, during and after any control attempts can provide valuable ecological information. In particular, it is important to understand how changes in the abundance of species influence ecosystem properties and processes which, in turn, will help guide management decisions. Ideally, this monitoring has to go beyond simple’ impacts on plant communities, involve cross-disciplinary teams of scientists and should incorporate many different taxa and their interactions. Monitoring design and data collection should be sophisticated enough to allow statistically sound data analysis. The available information will be paramount in (1) developing new political and scientific guidelines in invasive species management, (2) helping resolve potential conflicts of interest and (3) helping change public attitudes regarding growth, sale, and control of non-indigenous species.

Biological Control of Purple LoosestrifeA case for using insects as control agents, after rigorous screening, and for integrating release strategies with research

P urple loosestrife (Lythrum salicaria L.) is an exotic wetland perennial responsible for the degradation of many prime wetland habitats throughout the temperate regions of the United States and Canada. Large, monotypic stands reduce the biotic diversity of wetland systems by replacing native plant species (Stuckey 1980) and thereby eliminating the natural foods and cover essential to many wetland wildlife inhabitants, including waterfowl (Friesen 1966, Rawinski and Malecki 1984, Smith 1964). In North America, this plant is a classic example of an introduced species whose distribution and spread has been enhanced by the absence of natural enemies and the disturbance of natural systems, pri-

Mapping where ecologists work: biases in the global distribution of terrestrial ecological observations

Although the geographical context of ecological observations shapes ecological theory, the global distribution of ecological studies has never been analyzed. Here, we document the global distribution and context (protected status, biome, anthrome, and net primary productivity) of 2573 terrestrial study sites reported in recent publications (2004–2009) of 10 highly cited ecology journals. We find evidence of several geographical biases, including overrepresentation of protected areas, temperate deciduous woodlands, and wealthy countries. Even within densely settled or agricultural regions, ecologists tend to study “natural” fragments. Such biases in trendsetting journals may limit the scalability of ecological theory and hinder conservation efforts in the 75% of the terrestrial world where humans live and work.

Impact and management of purple loosestrife (Lythrum salicaria) in North America

The invasion of non-indigenous plants is considered a primary threat to integrity and function of ecosystems. However, there is little quantitative or experimental evidence for ecosystem impacts of invasive species. Justifications for control are often based on potential, but not presently realized, recognized or quantified, negative impacts. Should lack of scientific certainty about impacts of non-indigenous species result in postponing measures to prevent degradation? Recently, management of purple loosestrife (Lythrum salicaria), has been criticized for (1) lack of evidence demonstrating negative impacts of L. salicaria, and (2) management using biocontrol for lack of evidence documenting the failure of conventional control methods. Although little quantitative evidence on negative impacts on native wetland biota and wetland function was available at the onset of the control program in 1985, recent work has demonstrated that the invasion of purple loosestrife into North American freshwater wetlands alters decomposition rates and nutrient cycling, leads to reductions in wetland plant diversity, reduces pollination and seed output of the native Lythrum alatum, and reduces habitat suitability for specialized wetland bird species such as black terns, least bitterns, pied-billed grebes, and marsh wrens. Conventional methods (physical, mechanical or chemical), have continuously failed to curb the spread of purple loosestrife or to provide satisfactory control. Although a number of generalist insect and bird species utilize purple loosestrife, wetland habitat specialists are excluded by encroachment of L. salicaria. We conclude that (1) negative ecosystem impacts of purple loosestrife in North America justify control of the species and that (2) detrimental effects of purple loosestrife on wetland systems and biota and the potential benefits of control outweigh potential risks associated with the introduction of biocontrol agents. Long-term experiments and monitoring programs that are in place will evaluate the impact of these insects on purple loosestrife, on wetland plant succession and other wetland biota.

Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

Identification of factors that drive changes in plant community structure and contribute to decline and endangerment of native plant species is essential to the development of appropriate management strategies. Introduced species are assumed to be driving causes of shifts in native plant communities, but unequivocal evidence supporting this view is frequently lacking. We measured native vegetation, non-native earthworm biomass, and leaf-litter volume in 15 forests in the presence and absence of 3 non-native plant species (Microstegium vimineum, Alliaria petiolata, Berberis thunbergii) to assess the general impact of non-native plant and earthworm invasions on native plant communities in northeastern United States. Non-native plant cover was positively correlated with total native plant cover and non-native earthworm biomass. Earthworm biomass was negatively associated with cover of native woody and most herbaceous plants and with litter volume. Graminoid cover was positively associated with non-native earthworm biomass and non-native plant cover. These earthworm-associated responses were detected at all sites despite differences in earthworm species and abundance, composition of the native plant community, identity of invasive plant species, and geographic region. These patterns suggest earthworm invasion, rather than non-native plant invasion, is the driving force behind changes in forest plant communities in northeastern North America, including declines in native plant species, and earthworm invasions appear to facilitate plant invasions in these forests. Thus, a focus on management of invasive plant species may be insufficient to protect northeastern forest understory species.

Demographic models inform selection of biocontrol agents for garlic mustard (Alliaria petiolata).

Nonindigenous invasive plants pose a major threat to natural communities worldwide. Biological control of weeds via selected introduction of their natural enemies can affect control over large spatial areas but also risk nontarget effects. To maximize effectiveness while minimizing risk, weed biocontrol programs should introduce the minimum number of host-specific natural enemies necessary to control an invasive nonindigenous plant. We used elasticity analysis of a matrix model to help inform biocontrol agent selection for garlic mustard (Alliaria petiolata (M. Bieb.) Cavara and Grande). The Eurasian biennial A. petiolata is considered one of the most problematic invaders of temperate forests in North America. Four weevil species in the genus Ceutorhynchus (Coleoptera: Curculionidae) are currently considered potential biocontrol agents. These species attack rosettes (C. scrobicollis), stems (C. roberti, C. alliariae), and seeds (C. constrictus) of A. petiolata. Elasticity analyses using A. petiolata demographic parameters from North America indicated that changes in the rosette-to-flowering-plant transition and changes in fecundity consistently had the greatest impact on population growth rate. These results suggest that attack by the rosette-feeder C. scrobicollis, which reduces overwintering survival, and seed or stem feeders that reduce seed output should be particularly effective. Model outcomes differed greatly as A. petiolata demographic parameters were varied within ranges observed in North America, indicating that successful control of A. petiolata populations may occur under some, but not all, conditions. Using these a priori analyses we predict: (1) rosette mortality and reduction of seed output will be the most important factors determining A. petiolata demography; (2) the root-crown feeder C. scrobicollis will have the most significant impact on A. petiolata demography; (3) releases of single control agents are unlikely to control A. petiolata across its full range of demographic variability; (4) combinations of agents that simultaneously reduce rosette survival and seed production will be required to suppress the most vigorous A. petiolata populations. These predictions can be tested using established long-term monitoring sites coupled with a designed release program. If demographic models can successfully predict biocontrol agent impact on invasive plant populations, a continued dialogue and collaboration between empirical and theoretical approaches may be the key to the development of successful biocontrol tactics for plant invaders in the future.

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.

Invasive Plant and Experimental Venue Affect Tadpole Performance

Introductions of non-native predators and competitors appear to contribute to worldwide amphibian declines; however, potential negative impacts of invasive plants on habitat quality and amphibian populations have not been examined. Loss of diversity and alterations in ecosystem function associated with plant invasions may disrupt food webs, potentially leading to further declines of already threatened amphibian populations. We used a combination of small bins, mesocosms, and field experiments to examine the impacts of Eurasian purple loosestrife (Lythrum salicaria) replacing native cattails (Typha latifolia) in North American freshwater wetlands on survival, developmental rate, and diet (freshwater algae) of American toad (Bufo americanus) tadpoles. Tadpoles developed slower in L. salicaria compared to tadpoles developing in T. latifolia. This effect was consistent across experimental venues, although mesocosms showed this effect only in the second year of our study. Survival and development rates were always more variable in purple loosestrife than in cattail. In bins, tadpoles showed significantly reduced survival when raised in purple loosestrife extract and addition of leaf litter exacerbated this negative effect. Tadpole survival rates in mesocosms and field cages were not significantly different between plant species, most likely an effect of high variability among replicates. We suspect a combination of direct toxicity of high tannin concentrations in L. salicaria leaves and their indirect negative impacts on aquatic food webs are responsible for these results. Tadpole gut analyses revealed differences in algal communities among venues and between L. salicaria and T. latifolia suggesting that alterations in tadpole food quality and quantity contribute to the observed reduced tadpole performance. The replacement of native wetland plant species by L. salicaria does not represent a simple exchange of ecological equivalents and the function of invaded habitats for native species has clearly changed. While we were investigating only a single amphibian species, our results suggest that the impact of L. salicaria on ecosystem processes and aquatic food webs may be more general and likely to negatively affect other wetland species. The threats non-indigenous plants represent for amphibian populations and food webs may be underestimated, and warrant further investigation.

An evaluation of mechanisms preventing growth and survival of two native species in invasive Bohemian knotweed (Fallopia ×bohemica, Polygonaceae)

Restoring native diversity to habitats dominated by invasive plants requires improved understanding of mechanisms that allow introduced plants to retain dominance. We used a factorial transplant experiment to assess whether light limitation, nutrient limitation, or allelopathic interference by Fallopia ×bohemica reduces growth or survival of Eupatorium perfoliatum or Acer saccharinum. Increased light improved A. saccharinum performance but had no effect on E. perfoliatum growth. Increased light had no effect on A. saccharinum survival but improved E. perfoliatum survival. Activated carbon addition had short-term benefits allowing E. perfoliatum plants to grow for 4 wk and increased A. saccharinum leaf width but not biomass. Nutrient addition had no beneficial effect on transplants. These results in combination with the outcome of a cutting experiment suggest that F. ×bohemica achieves competitive superiority primarily by limiting access to light. Species-specific effects and significant interaction effects particularly of light and activated carbon suggest additional mechanisms. Allelopathic interference or interaction with microbial soil organisms may contribute to the lack of native species in populations of F. ×bohemica. Restoration efforts should combine selective F. ×bohemica control with soil amendments (such as activated carbon) to assess their potential to facilitate return of native species to invaded habitats.

Coexistence of Two Leaf-Beetles in the Same Fundamental Niche. Distribution, Adult Phenology and Oviposition

This study investigated the coexistence of two leaf-beetles, Galerucella calmariensis and G. pusilla, on purple loosestrife (Lythrum salicaria). G. pusilla and G. calmariensis share their distribution on a continental, regional, and local scale. Models of stable coexistence of two competitors assume access of inferior species to resources or refuges that are temporarily or spatially unexploited by the superior species. Field and laboratory studies indicated that both species have inclusive fundamental niches on their host plant. Cytological studies and crossing experiments confirmed their status as true, reproductively isolated species. G. calmariensis and G. pusilla showed no differences in adult phenology, winter mortality, colonization pattern nor oviposition, thus no temporal variation in resource use. Adults aggregated at the same sites and on the same host plant individuals and their distributions are not independant. Aggregation or differences in resource use do not appear to play an important role in the coexistence of G. calmariensis and G. pusilla. This study indicated that both species have identical competitive abilities.