Betsy Von Holle

Positive interactions of nonindigenous species: invasional meltdown?

Study of interactions between pairs or larger groups of nonindigenous species has been subordinated in the literature to study of interactions between nonindigenous and native species. To the extent that interactions among introduced species are depicted at all, the emphasis has been on negative interactions, primarily resource competition and interference. However, a literature search reveals that introduced species frequently interact with one another and that facilitative interactions are at least as common as detrimental ones. The population significance of these interactions has rarely been determined, but a great variety of types of direct and indirect interactions among individuals of different nonindigenous species is observed, and many are plausibly believed to have consequences at the population level. In particular, mutualisms between plants and the animals that disperse and/or pollinate them and modification of habitat by both animals and plants seem common and often important in facilitating invasions. There is little evidence that interference among introduced species at levels currently observed significantly impedes further invasions, and synergistic interactions among invaders may well lead to accelerated impacts on native ecosystems – an invasional ‘meltdown’ process.

Loss of foundation species: consequences for the structure and dynamics of forested ecosystems

In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species-specific traits control ecosystem dynamics. Such foundation tree species are declining throughout the world due to introductions and outbreaks of pests and pathogens, selective removal of individual taxa, and over-harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environment on which a variety of other species depend; how this disrupts fundamental ecosystem processes, including rates of decomposition, nutrient fluxes, carbon sequestration, and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynamics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small perturbations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, anticipate the cascade of immediate, short- and long-term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management.

ECOLOGICAL RESISTANCE TO BIOLOGICAL INVASION OVERWHELMED BY PROPAGULE PRESSURE

Models and observational studies have sought patterns of predictability for invasion of natural areas by nonindigenous species, but with limited success. In a field experiment using forest understory plants, we jointly manipulated three hypothesized determinants of biological invasion outcome: resident diversity, physical disturbance and abiotic conditions, and propagule pressure. The foremost constraints on net habitat invasibility were the number of propagules that arrived at a site and naturally varying resident plant density. The physical environment (flooding regime) and the number of established resident species had negligible impact on habitat invasibility as compared to propagule pressure, despite manipulations that forced a significant reduction in resident richness, and a gradient in flooding from no flooding to annual flooding. This is the first experimental study to demonstrate the primacy of propagule pressure as a determinant of habitat invasibility in comparison with other candidate controlling factors.

Economic Impacts of Non-Native Forest Insects in the Continental United States

Reliable estimates of the impacts and costs of biological invasions are critical to developing credible management, trade and regulatory policies. Worldwide, forests and urban trees provide important ecosystem services as well as economic and social benefits, but are threatened by non-native insects. More than 450 non-native forest insects are established in the United States but estimates of broad-scale economic impacts associated with these species are largely unavailable. We developed a novel modeling approach that maximizes the use of available data, accounts for multiple sources of uncertainty, and provides cost estimates for three major feeding guilds of non-native forest insects. For each guild, we calculated the economic damages for five cost categories and we estimated the probability of future introductions of damaging pests. We found that costs are largely borne by homeowners and municipal governments. Wood- and phloem-boring insects are anticipated to cause the largest economic impacts by annually inducing nearly

Historical Accumulation of Nonindigenous Forest Pests in the Continental United States

1.7 billion in local government expenditures and approximately

Economic Impacts of Invasive Species in Forests Past, Present, and Future

830 million in lost residential property values. Given observations of new species, there is a 32% chance that another highly destructive borer species will invade the U.S. in the next 10 years. Our damage estimates provide a crucial but previously missing component of cost-benefit analyses to evaluate policies and management options intended to reduce species introductions. The modeling approach we developed is highly flexible and could be similarly employed to estimate damages in other countries or natural resource sectors.

The importance of biological inertia in plant community resistance to invasion

Nonindigenous forest insects and pathogens affect a range of ecosystems, industries, and property owners in the United States. Evaluating temporal patterns in the accumulation of these nonindigenous forest pests can inform regulatory and policy decisions. We compiled a comprehensive species list to assess the accumulation rates of nonindigenous forest insects and pathogens established in the United States. More than 450 nonindigenous insects and at least 16 pathogens have colonized forest and urban trees since European settlement. Approximately 2.5 established nonindigenous forest insects per year were detected in the United States between 1860 and 2006. At least 14% of these insects and all 16 pathogens have caused notable damage to trees. Although sap feeders and foliage feeders dominated the comprehensive list, phloem- and wood-boring insects and foliage feeders were often more damaging than expected. Detections of insects that feed on phloem or wood have increased markedly in recent years.

How many, and which, plants will invade natural areas?

Biological invasions by nonnative species are a by‐product of economic activities, with the vast majority of nonnative species introduced by trade and transport of products and people. Although most introduced species are relatively innocuous, a few species ultimately cause irreversible economic and ecological impacts, such as the chestnut blight that functionally eradicated the American chestnut across eastern North America. Assessments of the economic costs and losses induced by nonnative forest pests are required for policy development and need to adequately account for all of the economic impacts induced by rare, highly damaging pests. To date, countrywide economic evaluations of forest‐invasive species have proceeded by multiplying a unit value (price) by a physical quantity (volume of forest products damaged) to arrive at aggregate estimates of economic impacts. This approach is inadequate for policy development because (1) it ignores the dynamic impacts of biological invasions on the evolution of prices, quantities, and market behavior, and (2) it fails to account for the loss in the economic value of nonmarket ecosystem services, such as landscape aesthetics, outdoor recreation, and the knowledge that healthy forest ecosystems exist. A review of the literature leads one to anticipate that the greatest economic impacts of invasive species in forests are due to the loss of nonmarket values. We proposed that new methods for evaluating aggregate economic damages from forest‐invasive species need to be developed that quantify market and nonmarket impacts at microscales that are then extended using spatially explicit models to provide aggregate estimates of impacts. Finally, policies that shift the burden of economic impacts from taxpayers and forest landowners onto parties responsible for introducing or spreading invasives, whether through the imposition of tariffs on products suspected of imposing unacceptable risks on native forest ecosystems or by requiring standards on the processing of trade products before they cross international boundaries, may be most effective at reducing their impacts.

Climatic variability leads to later seasonal flowering of Floridian plants.

Abstract Insights into the ecology of historic invasions by introduced species can be gained by studying long-term patterns of invasions by native species. In this paper, we review literature in palaeo-ecology, forest-stand simulation modelling, and historical studies of plant species invasions to illustrate the relevance of biological inertia in plant communities to invasion ecology. Resistance to invasion occurs in part because of environmental, demographic, and biotic factors influencing the arrival and establishment of invading species. We propose that biological inertia within the resident community is a fourth component of resistance to invasion, because of the lag time inherent in eliminating resident species and perhaps their traces after environmental conditions become suitable for invasion by immigrating species. Whether or not an introduced species invades can be conditioned by the presence of the pre-existing community (and/or its legacy) in addition to the other biotic and abiotic factors.

Invasive Plant Species and the Ornamental Horticulture Industry

Of established nonindigenous plant species in California, Florida, and Tennessee, 5.8%, 9.7%, and 13.4%, respectively, invade natural areas according to designations tabulated by state Exotic Pest Plant Councils. Only Florida accords strictly with the tens rule, though California and Tennessee fall within the range loosely viewed as obeying the rule. The species that invaded natural areas in each state were likely, if they invaded either of the other states at all, to have invaded natural areas there. There was a detectable but inconsistent tendency for species that invade natural areas to come from particular families. At the genus level in California and Florida, and the family level in California, there was also a tendency for natural area invaders to come from taxa that were not represented in the native flora. All three of the above patterns deserve further studies to determine management implications. Only the first (that natural area invaders of one state are likely to invade natural areas if they invade another state) seems firm enough from our data to suggest actions on the part of managers.