Showing papers in "Biological Invasions in 1999"

Positive interactions of nonindigenous species: invasional meltdown?

Abstract: 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.

Impact: Toward a Framework for Understanding the Ecological Effects of Invaders

Abstract: Although ecologists commonly talk about the impacts of nonindigenous species, little formal attention has been given to defining what we mean by impact, or connecting ecological theory with particular measures of impact. The resulting lack of generalizations regarding invasion impacts is more than an academic problem; we need to be able to distinguish invaders with minor effects from those with large effects in order to prioritize management efforts. This paper focuses on defining, evaluating, and comparing a variety of measures of impact drawn from empirical examples and theoretical reasoning. We begin by arguing that the total impact of an invader includes three fundamental dimensions: range, abundance, and the per-capita or per-biomass effect of the invader. Then we summarize previous approaches to measuring impact at different organizational levels, and suggest some new approaches. Reviewing mathematical models of impact, we argue that theoretical studies using community assembly models could act as a basis for better empirical studies and monitoring programs, as well as provide a clearer understanding of the relationship among different types of impact. We then discuss some of the particular challenges that come from the need to prioritize invasive species in a management or policy context. We end with recommendations about how the field of invasion biology might proceed in order to build a general framework for understanding and predicting impacts. In particular, we advocate studies designed to explore the correlations among different measures: Are the results of complex multivariate methods adequately captured by simple composite metrics such as species richness? How well are impacts on native populations correlated with impacts on ecosystem functions? Are there useful bioindicators for invasion impacts? To what extent does the impact of an invasive species depend on the system in which it is measured? Three approaches would provide new insights in this line of inquiry: (1) studies that measure impacts at multiple scales and multiple levels of organization, (2) studies that synthesize currently available data on different response variables, and (3) models designed to guide empirical work and explore generalities.

Behavioral and genetic differentiation between native and introduced populations of the Argentine ant.

Abstract: In this paper, we examine the hypothesis that reduced intraspecific aggression underlies the competitive prowess of Argentine ants in their introduced range. Specifically, we test three predictions of this hypothesis by comparing the genetic diversity, behavior, and ecology of Argentine ants in their native range to introduced populations. Differences between native and introduced populations of Argentine ants were consistent with our predictions. Introduced populations of the Argentine ant appear to have experienced a population bottleneck at the time of

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

Abstract: 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.

When to Ignore Advice: Invasion Predictions and Decision Theory

Abstract: Organisms generally become pests at a low rate. As a consequence of this low ‘base-rate probability’, the large majority of organisms rejected in any random sample of potential introductions would probably be harmless, despite the fairly high accuracy of some recently proposed risk assessment systems for exotic introductions. Here we distinguish between a system's accuracy (the proportion of a group of known pest species that would be correctly identified as pests) and reliability (the rate of false positives and false negatives produced once the base-rate is taken into account). We next adapt a decision theory analysis of earthquake prediction to explore when we would be best advised to ignore the recommendations of a screening system for exotic introductions. In one scenario, we show that a pest risk assessment system with an accuracy of 85% would be better ignored, unless the damage caused by introducing a pest is eight times or more that caused by not introducing a harmless organism that is potentially useful. Furthermore, because of the base-rate effect, in certain situations it may be more efficient to focus on identifying potential invaders from amongst already naturalized species than from amongst species at the importation stage.

Experimental Invasions Using Biological Control Introductions: The Influence of Release Size on the Chance of Population Establishment

Abstract: Introductions of biological control organisms offer a unique opportunity to experimentally study the process of invasion by exotic species. I used two chrysomelid beetles, Galerucella calmariensis and Galerucella pusilla, which are currently being introduced into North America for the biological control of purple loosestrife (Lythrum salicaria), to determine how the initial size of a release affects the probability that the introduced population grows and persists. I released both species into stands of their host plant at 36 sites scattered throughout central New York State using four release sizes: 20, 60, 180, and 540. I returned to these sites over the next 3 years to census the populations. For both species, the probability of population establishment increased with release size. Population growth rates also depended positively on release size. The implication from these results is that the demographic factors whose influence depends on population size or density such as demographic stochasticity, Allee effects, and genetics play important roles in the establishment of invading populations. A second set of releases was used to determine if it was at all possible for a single gravid female to found a population. Out of twenty individual females released, one female (a G. calmariensis) founded a population that persisted until the end of the study (3 generations).

The Disturbed Resource-Flux Invasion Matrix: A New Framework for Patterns of Plant Invasion

Abstract: Attempts to classify certain habitats as vulnerable to invasion or plant traits as invasive have met with limited success and applicability. Clearly, not all plant invaders are able to exploit all habitats and not all habitats are equally susceptible to invasion. Here we argue that it is critical for a successful model for invasions to incorporate both environmental and species traits and present just such a framework. Although disturbance has been targeted as a crucial event which renders habitats vulnerable to invasion, disturbances are often integral parts of ecosystems (e.g. floods, tree-falls, fire, etc.) and are not always associated with invasion events. We argue that disturbances that are associated with invasions alter historical patterns of turnover, or flux, of resources in an ecosystem. Given this perspective on the relationship between invasions and disturbances, and the need to integrate species traits with those of invaded ecosystems, we have developed an approach to characterize plant invasion patterns that we call the ‘Disturbed Resource-Flux Invasion Matrix’ or DRIM. This is a 16-cell matrix that classifies habitats by the quality of changes in physical and chemical resource flux either increasing or decreasing flux relative to historical patterns. Within each matrix cell, it is then possible to apply basic ecological principles to target species traits that can facilitate successful invasion of habitats experiencing that particular kind of disturbance. We present examples from the literature of how habitats and species can be classified according to the DRIM, and demonstrate the application of this theoretical model.

The Invasion Ecology of Japanese Barberry (Berberis thunbergii) in the New England Landscape

Abstract: Japanese barberry (Berberis thunbergii) has been characterized as one of the most widely known and planted exotic shrubs in the United States. It was first introduced to the US in the late 1800s. By 1920 the planting of Japanese barberry was encouraged as an ornamental shrub replacing the common barberry (Berberis vulgaris). Japanese barberry began spreading from cultivation in suburban and selected rural retreats by the 1920s, and had dispersed rapidly throughout the northeast by the 1960s. By the 1970s it was recognized as a problematic invasive in the northeast. It is readily dispersed primarily by birds. Fruit production varies with light level, but even under very low light levels (≤4% full sun) some seeds are produced. Fruits are dispersed in late fall through late winter. Seed dispersal curves are highly leptokurtic; most seedling are found under or adjacent to adults, but a small number may be found tens of meters from the nearest adult. Japanese barberry thrives under a broad range of light and soil moisture conditions. Significant variation in stem growth can be explained as a function of light level. Even at less than 1% full sun, some positive stem growth can occur. Survival is quite high at intermediate to high light levels, and only under the lowest light levels (<1% full sun) does survival drop significantly. Biomass of Japanese barberry in field plots can be largely explained as a function of light availability and soil moisture. The biomass of co-occurring species is suppressed by Japanese barberry, and recovery is slow in the first year following Japanese barberry removal except under high light levels. Glyphosate (Roundup) applied in early spring at first leaf out, when little else is in leaf, provides an effective means of eradicating Japanese barberry populations.

Differences in earthworm densities and nitrogen dynamics in soils under exotic and native plant species.

Abstract: Previous studies of the invasion of two exotic plants – Berberis thunbergii and Microstegium vimineum – in hardwood forests of New Jersey have shown a significant increase of pH in soils under the invasive plants as compared with soils from under native shrubs (Vaccinium spp). We present a further investigation of soil properties under the exotic plants in question. We measured the densities of earthworms in the soil under the two exotics and the native shrubs in three parks in New Jersey. In the same populations we also measured the extractable ammonium and nitrate in the top 5 cm of the soil, as well as the respiration of the soils and the potential rates of mineralization (aerobic lab incubation). In addition, we measured the nitrate reductase activity in leaves of the two exotic plants and several native shrubs and trees. Although there were differences between parks, we observed significantly higher earthworm densities in the soil under the exotic species. The worms were all European species. Soil pH, available nitrate and net potential nitrification were significantly higher in soils under the two exotic species. In contrast, total soil C and N and net ammonification were significantly higher under native vegetation. Nitrate reductase activities were much higher in the leaves of exotic plants than in the leaves of native shrubs and trees. Changes in soil properties, especially the change in nitrogen cycling, associated with the invasion of these two plant species may permit the invasion of other weedy or exotic species. Our results also suggest that even if the two exotic species were removed, the restoration of the native flora might be inhibited by the high nitrate concentrations in the soil.

Invasion Pattern of Herb Garlic Mustard (Alliaria petiolata) in High Quality Forests

Abstract: The invasion of non-indigenous plant species poses a severe threat to native plant communities. Garlic mustard (Alliaria petiolata) is a naturalized European biennial herb that has spread rapidly through the eastern US and adjacent Canada. To determine garlic mustard rate of spread, eleven permanent plots (50×25 m) were located in seven high quality (relatively undisturbed) forests in the early stages of invasion. Garlic mustard presence was recorded within six 50×2 m permanent belt transects, and density and percent cover by age class were recorded in 36 permanent 1 m2 quadrats, between 1989 and 1992, and again in 1997. Garlic mustard spread at an average rate of 5.4 m per year between 1989 and 1992, in all plots combined. Within individual plots rate of spread varied substantially, with location of the front increasing up to 36 m and decreasing as much as 18 m between years. While the front alternately advanced and retreated, over time garlic mustard consistently advanced through all forests. Rate of spread was influenced by establishment of satellite populations, and disturbance (wind-throw and flooding). The pattern of spread within plots was one of a ragged advancing front, supplemented by establishment of satellite populations 6–40 m distant from the front, which then coalesced with the main population. Garlic mustard presence between 1989 and 1997 increased significantly within all plots, and in each age class within each plot. The greatest increases occurred in plots where this plant was initially rarest. Garlic mustard cover and density varied nonsignificantly during the same time period. These results indicate that after garlic mustard invades a forest it becomes a permanent part of the community, annually increasing in presence but fluctuating in cover and density. Garlic mustard maintains a low profile under low disturbance conditions, but increases rapidly with periodic disturbance. This study monitored garlic mustard invasion in high quality relatively undisturbed forests, and may underestimate the rate of spread in low quality highly disturbed forests.

Invasion pressure to a ballast-flooded estuary and an assessment of inoculant survival

Abstract: The relationships between invasion pressure, post-transport inoculant survival, and regional susceptibility to invasion are poorly understood. In marine ecosystems, the movement and release of ballast water from ocean-going ships provides a model system by which to examine the interplay among these factors. One of the largest estuaries in North America, the Chesapeake Bay, receives tremendous amounts of foreign ballast water annually and thus should be at high invasion risk. To date, however, few introductions in Chesapeake Bay have been attributed to ballast release. To understand better the dynamics of this invasion process, we (1) characterized and quantified the biota arriving to Chesapeake Bay in foreign ballast water, (2) compared temperatures and salinities of ballast water and harbor water in upper Chesapeake Bay, and (3) tested experimentally survival of organisms collected from ballast water in temperatures and salinities characteristic of the region. From 1993 to 1994, we sampled planktonic and benthic organisms from 60 foreign vessels arriving to Chesapeake Bay. Our data show that the estuary is being inoculated by a diverse assemblage of aquatic organisms from around the world. Furthermore, the short transit time (≤15 d) for most vessels ensured that substantial numbers of larval and post-larval organisms were being deballasted alive. Most of the ballast water discharged into the upper Chesapeake Bay, however, was significantly higher in salinity (>20‰) than that of the receiving harbor. In laboratory tolerance experiments, ballast water organisms perished under such conditions. Thus, a mismatch in physical conditions between donor and receiver regions may explain the dearth of invasions in the upper Bay. It is likely that the lower Chesapeake Bay, which is more saline, remains at higher risk to ballast water invasion. Recognition of such intraregional differences should allow more focused predictions for monitoring and management.

Introduction of Nonindigenous Aquatic Vascular Plants in Southern New England: A Historical Perspective

Abstract: Aquatic plants comprise few species worldwide, yet introductions of nonindigenous hydrophytes represent some of the most severe examples of biological invasions. Often innocuous in their indigenous regions, many aquatic plant species have caused extreme ecological and economic consequences when introduced into nonindigenous habitats. Typically, aquatic plant invasions are unnoticed or overlooked until they are perceived as problematic. By then, plants are virtually impossible to eradicate and negative ecological impacts caused by their spread into natural communities are irreparable. We present criteria to facilitate decisions whether a species should be characterized as nonindigenous or invasive. Historical data are used to clarify methods of introduction, avenues and means of dispersal, and extent of invasiveness of the following aquatic plants in southern New England: Acorus calamus, Butomus umbellatus, Cabomba caroliniana, Callitriche stagnalis, Egeria densa, Hydrilla verticillata, Limnobium spongia, Marsilea quadrifolia, Myriophyllum aquaticum, Myriophyllum heterophyllum, Myriophyllum spicatum, Najas minor, Najas guadalupensis, Nasturtium officinale, Nymphoides peltata, Potamogeton crispus, Trapa natans and Veronica beccabunga.

Norway Maple (Acer platanoides) Invasion of a Natural Forest Stand: Understory Consequence and Regeneration Pattern

Abstract: Norway maple (Acer platanoidesis) is invasive in a natural stand in suburban Ithaca, NY. To determine the understory pattern and consequences of a Norway maple invasion, I compared density and species richness under Norway maples and sugar maples (Acer saccharum). Mean sapling density was significantly lower (P<0.0027) under Norway maples (3.64/100 m2±1.6 SE) than under sugar maples (19.4/100 m2±4.4 SE). Mean sapling species richness was significantly lower (P<0.0018) under Norway maples (0.7/32 m2±0.18 SE) than under sugar maples (2.6/32 m2±0.48 SE). Likewise, Norway maple regeneration is more frequent under sugar maples than sugar maple regeneration: 57% of sugar maple plots had Norway maple saplings while 0% of Norway maple plots had sugar maple saplings. Two significant plot effects were found for presence–absence: Norway maple saplings grow under Norway maples with a significantly lower frequency (P<0.03) than under sugar maples; sugar maple saplings grow under Norway maples with a significantly lower frequency (P<0.000) than under sugar maples. Across the site, Norway maple saplings were the most abundant (29 saplings for 480 m2). The success of Norway maple regeneration and the reductions in total stem density beneath Norway maples is most likely the result of its strong competitive abilities, notably its high shade tolerance and abundant seed crops.

Reshaping the Ecology of Invading Populations of Hemlock Woolly Adelgid, Adelges tsugae (Homoptera: Adelgidae), in Eastern North America

Abstract: Hemlock woolly adelgid, Adelges tsugae Annand (Homoptera: Adelgidae), is native to Japan where it is an innocuous inhabitant of Tsuga diversifolia Masters and T. sieboldii Carriere throughout their natural growing areas. Native adelgid populations are regulated by host resistance and natural enemies, in particular the oribatid mite, Diapterobates humeralis (Hermann) and the coccinellid beetle, Pseudoscymnus tsugae Sasaji and McClure. Invading populations of A. tsugae in western North America on T. heterophylla Sargent and T. mertensiana Carriere are mainly regulated by host resistance. However, invading populations in eastern North America attain damaging levels on T. canadensis (L.) Carriere and T. caroliniana Engelmann and are regulated mainly by weather and negative density-dependent feedback mechanisms related to host deterioration. Although A. tsugae is only passively dispersed by wind, birds, forest-dwelling mammals and humans, it is spreading at an alarming rate and is sufficiently cold hardy to threaten the existence of the two eastern hemlock species throughout their natural ranges. The current hope for suppressing invading populations of hemlock woolly adelgid in eastern North America lies with the exotic predator, P. tsugae. Extensive laboratory studies of the biology and predatory ability of P. tsugae revealed that it feeds on all life stages of its prey, that its multivoltine life cycle is well synchronized with that of the adelgid, and that it has great potential for biological control. We have reared and released nearly 130,000 adults of P. tsugae in forests in Connecticut, New Jersey and Virginia during the past four years. P. tsugae has reproduced, dispersed, overwintered and reduced densities of hemlock woolly adelgid by 47–88% in only five months on release branches at these sites. Current studies are investigating the long-term ability of P. tsugae to regulate invading populations of A. tsugae in eastern North America.

A Paleoecological Assessment of Phragmites australis in New England Tidal Marshes: Changes in Plant Community Structure During the Last Few Millennia

Abstract: Although Phragmites has been an upper border tidal marsh species for thousands of years, it is only recently (within the last century or so) that the distribution of this plant within the coastal marsh community has become prominent. Prior to approximately 100 years ago, Phragmites was an upper border/brackish marsh co-dominant in many marsh systems. Occurrence of this species varied between associations of sedges, Typha, forbs and a variety of woody shrubs. Paleoreconstructions rarely show the presence of a Phragmites monoculture or early associations with salt marsh species. However, since the turn of this century (and perhaps as early as the middle of the last century) the distribution of Phragmites has changed substantially. Today, this plant often forms dense monocultures and is commonly found in association with Spartina grasses. The results of this paleoecological investigation show that the changes that have been observed in Phragmites communities during the last 100 years are not part of the long-term cycle of development in these systems and are new to the landscape.

Habitat Invasibility and Dominance by Alien Annual Plants in the Western Mojave Desert

Abstract: Patterns of habitat invasibility and alien dominance, respectively measured as species richness and biomass of alien annual plants, were evaluated in association with four habitat factors at the Desert Tortoise Research Natural Area (DTNA) in the western Mojave Desert, USA. Habitat factors varied in levels of disturbance outside (high) and inside (low) the DTNA, and in levels of soil nutrients in washlet (high) and hummock (low) topographic positions, in Larrea-north (high), Larrea-south (medium), and interspace (low) microhabitats near creosote bushes (Larrea tridentata), and during 1995 when rainfall was 207% (high) and 1994 when rainfall was 52% (low) of the long-term average. Dominant alien plants included the annual grasses Bromus rubens, Bromus trinii, and Schismus spp., and the forb Erodium cicutarium. Species richness and dominance of alien annual plants were slightly higher where disturbance was high, and much higher where soil nutrients were high. B. rubens and B. trinii were most dominant in washlets and in the Larrea-north microhabitats during both years. These two species evolved in mesic ecosystems, and appeared to be particularly limited by soil nutrients at this site. Schismus spp. and E. cicutarium were also most dominant in washlets, but their dominance varied between interspaces in 1994 and the Larrea-south microhabitat in 1995. Monitoring to detect the invasion of new annual plants should focus on regions of high rainfall and nitrogen deposition and on washes and beneath-canopy microhabitats. The ecological range of each alien species should be evaluated separately, because their evolutionary origins may greatly affect their patterns of invasion and dominance in the Mojave Desert.

The Invasive Woody Weed Ligustrum robustum subsp. walkeri Threatens Native Forests on La Réunion

Abstract: One of the last primitive island ecosystems in the Indian Ocean has been invaded since 1969 by the Sri Lankan privet, Ligustrum robustum. L. robustum is still spreading in the forests of La Reunion Island, where only 30% of the original vegetation remains, but where 98% of the primary native vegetation of the Mascarene Islands still exists. On Mauritius, where L. robustum was introduced about 1895, it now forms dense, impenetrable thickets, and its presence has been correlated with the inability of native vegetation to re-establish. We assessed the potential impacts of L. robustum invasion on the native ecosystems of La Reunion and identified the factors of invasibility. We determined the degree of invasion in 12 plots of 156 m2 and followed native flora and privet recruitment for 3 years in 12 seedling plots of 39 m2. The data show that monocultural L. robustum stands now exist in human-disturbed primary forest patches (3.3 individuals/m2 and 80% of total individuals) and high seedling densities (0.3–0.6/m2) occur in the least disturbed patches. L. robustum's rapid growth, high shade tolerance and seed production, bird-assisted seed dispersal and high seedling recruitment contribute to its invasiveness in intact forests. The conservation of the original ecosystems of La Reunion depends on the setting up of a long-term and immediate global control strategy.

Establishment and Control of Hay-scented Fern: A Native Invasive Species

Abstract: Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m2 and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.

Species composition and inter-annual dynamics of a freshwater tidal plant community following removal of the invasive grass, Phragmites australis

Abstract: We document the regeneration of native freshwater wetland plant assemblages following removal of the common reed, Phragmites australis (Cav.) Trin. ex Steudel from two sites at Chapman Pond, East Haddam, Connecticut, USA. We gathered field data on composition of the vegetation 1 year before and for each of the 3 years after the removal in fall 1995/spring 1996 of Phragmites by two slightly different methods (hand-removal and herbiciding in one area, mowing/mulching and herbiciding in another). An area where Phragmites was left intact was similarly monitored. Our goals for this monitoring were: (1) to document plant species composition and richness before and after Phragmites removal and (2) to examine temporal and spatial variability in patterns of plant recruitment. Phragmites declined in both density and extent in both plots where removal treatments were applied. Richness, evenness, and density of non-Phragmites species increased steadily from 1996 to 1997 in all removal and intact plots. However, the species composition of the removal plots was richer than that of the intact plot, and more closely resembled that of comparable, uninvaded freshwater tidal wetlands. Rates of recovery of species richness in the removal plots declined from 1997 to 1998, potentially reflecting saturation of available colonization space, or the return of Phragmites. Phragmites has expanded its range in both of the removal plots since 1997. A model of its colonization indicates that Phragmites occupies space through localized proliferation of dense rhizomes rather than diffusely foraging with long tillers. Vigilance in monitoring is needed to document the spread of invasives, to evaluate the multi-faceted ecological effects of eradication efforts on both the invader and the regenerating community, and develop strategies for preventing re-invasion.

Structure and Dynamics of Populations of Japanese Barberry (Berberis Thunbergii DC.) in Deciduous Forests of New Jersey

Abstract: Japanese barberry, Berberis thunbergii DC., has become a prominent exotic species in deciduous forests throughout the eastern and midwestern US. Populations range from small plants occurring at low densities to dense, impenetrable thickets of plants with up to 40 stems/individual. A study was undertaken at Morristown National Historical Park in New Jersey to document plant densities, plant size, recruitment through vegetative processes of new shoot initiation and clonal spread and recruitment from seedling establishment, and mortality of stems and plants. Nearly 2000 shoots on 370 plants were individually marked and followed for two growing seasons, and over 1000 seedlings were also individually marked and followed. Populations vary much more in total shoots/area than they do in plant individuals/area, or in mean plant size (shoots/plant), as even the sparse populations have a few large individuals. Shoot mortality is less than new shoot initiation, but most plants do not change in size or change by small numbers of stems. However, the number of new shoots per plant increases as plant size increases. Once plants have three stems, they suffer little or no mortality. Seedling establishment is proportional to the density of shoots, so that as plants grow in size, local recruitment from seed increases. Large numbers of seedlings, and a survival rate of 10%, combine to make seedling recruitment a major component of population increase. The combination of multiple forms of vegetative and seed-based population growth, and the very low rates of plant mortality due to the multi-stemmed growth form explains the ability of this invasive species to rapidly produce dense, persistent populations.

Potential Impacts of a Western Pacific Grapsid Crab on Intertidal Communities of the Northwestern Atlantic Ocean

Abstract: Population density and size distribution, salinity tolerance, and feeding activity were examined in a western Pacific grapsid crab, Hemigrapsus sanguineus, that was recently introduced to the mid-Atlantic coast of North America. Seasonal abundance on a boulder/cobble shore (Crane Neck Pt.) in central Long Island Sound, New York, USA, during 1997–1998, ranged from 7 to 10 crabs m−2 averaged over the entire intertidal zone. Crabs occurred throughout the intertidal during summer and fall, but appeared to move from high to low elevations during winter. In laboratory experiments, H. sanguineus tolerated salinity down to 10 ppt for 7 d, but showed significant preference for 20 or 27 ppt over 10 ppt. The crabs readily consumed juvenile snails (Littorina littorea) and mussels (Mytilus edulis), as well as other common species of macroalgae and invertebrates occurring at Crane Neck Pt. High feeding rates and the ability to consume littorine snails up to 13 mm in height and mussels up to 20 mm in length suggest that this nonindigenous species has the potential to significantly affect the structure of rocky intertidal communities in the northwestern Atlantic Ocean; however, rigorous field studies are needed to accurately determine the impact of this recent introduction.

An Experimental Study of Competition Between the Native Sacramento Perch (Archoplites interruptus) and Introduced Bluegill (Lepomis macrochirus)

Abstract: The Sacramento perch (Archoplites interruptus), a sunfish (Centrarchidae) native only to the Central Valley of California, has been eliminated from most of its native range. To examine the role of interspecific competition in this decline, a series of experiments were conducted to assess the growth, aggressive behavior, and habitat use of Sacramento perch in the presence of bluegill (Lepomis macrochirus), an introduced centrarchid. The experiments indicate that (1) Sacramento perch gain less weight and show reduced growth when placed with bluegill, but that this interaction only occurs with food limitation, and is not affected by overall fish density; (2) Sacramento perch demonstrate less aggressive behavior than bluegill, but become more aggressive when they are conspicuously larger than bluegill; (3) Sacramento perch shift their habitat use in the presence of bluegill. Overall the results imply that Sacramento perch and bluegill exhibit interspecific competition where the mechanism of interaction is aggressive dominance by bluegill. It is suggested that long term persistence of Sacramento perch may require a habitat that is free of introduced centrarchid fishes, or one controlled by a naturally variable hydrological regime.

The Distribution of an Introduced Mollusc and its Role in the Long-term Demise of a Native Confamilial Species

Abstract: The estuarine mud snail, Batillaria attramentaria, was transported to the Pacific coast of North America with the Pacific oyster, Crassostrea gigas (C. gigas), imported from Japan in the early part of this century, and has proliferated in several bays where the oyster was introduced. Since the arrival of Batillaria there have been declines in populations of its native ecological equivalent, Cerithidea californica. This study documents the distribution of the exotic Batillaria throughout its entire introduced range, concentrating on the few bays in northern California where both snails exist sympatrically. Using dates of initial importation of C. gigas and dates of first documentation of Batillaria within a bay, I established the earliest possible date for the introduction of Batillaria in a particular area. In cases where Cerithidea also was, or had been, present within a bay, I calculated a range of time for either the continued coexistence of the two species or the time until local exclusion of the native. Density measurements of Cerithidea within these bays where the species co-occurred allowed comparison of present Cerithidea numbers to historical accounts. Results indicated that Batillaria is replacing Cerithidea in the northern marshes of California. This replacement of the northernmost Cerithidea populations is not only reducing Cerithidea's overall range, but also eliminating a race recently shown to be a genetically distinct from southern Cerithidea populations. Other studies that have demonstrated superior exploitative competitive ability by Batillaria provide a potential mechanism for this displacement pattern. Regardless, the results presented here indicate that the displacement process is slow, taking on average >50 years to complete. This study illustrates a gradual, but predictable process of exotic replacement of native species, and argues strongly against complacency toward invaders that may currently seem innocuous.

The Effects of Phragmites Removal on Nutrient Pools in a Freshwater Tidal Marsh Ecosystem

Abstract: Selected nitrogen and phosphorus pools in two freshwater tidal marsh ecosystems on the lower Connecticut River were measured relative to Phragmites, Typha and mixed native wetland plant cover types. For both the Chapman Pond Preserve and Chester Creek Marsh, significant differences were found between plant cover types in porewater ammonium and phosphate for some months during the 1998 growing season; porewater nitrate was always below the detection limit. At Chapman Pond, no significant differences were detected between Phragmites and Typha for plant tissue N concentrations. The standing stock of nitrogen was higher in Phragmites stands, however, owing to its greater aboveground biomass. No significant difference was found between plant cover types for total soil N at Chapman Pond, but KCl extractable ammonium was higher in the mixed cover type than Phragmites or Typha. The results of this study suggest that Phragmites is affecting nutrient pools in freshwater tidal marshes, a result that should be considered in future management design.

Using Satellite Images to Classify and Analyze the Health of Hemlock Forests Infested by the Hemlock Woolly Adelgid

Abstract: A method is described to classify stands of eastern hemlock by health condition, at the landscape level, using remote sensing. The hemlock woolly adelgid has been a major cause of hemlock decline in Connecticut since 1985, resulting in varying degrees of defoliation in the region. A 1985 Landsat Thematic Mapper (TM) image was classified to develop a base line of once healthy hemlock stands. Radiance normalization and non-hemlock masking techniques were used to pre-process a 1995 TM image. Several techniques were used to transform the 1995 TM image; each was followed by cluster analysis to separate hemlocks into four levels of tree vigor. We evaluated 600 trees at 150 sites across the study area using the USFS Crown Condition Rating Guide. These field data were used to measure the accuracy of various health classification techniques. The Modified Soil Adjusted Vegetation Index-2 (MSAVI2) transform provided the best overall accuracy, 82.1%, for classifying hemlock according to tree vigor. Non-parametric statistics were used to determine if there were any significant variations in distribution of hemlock pixels by health class in association with features in the landscape. Several features were found to be statistically significant at a confidence level of 0.001. These were aspect of slope, hydrology group (infiltration rate), depth to bedrock, soil order, drainage class (hydraulic conductivity), and surface texture.

Experimental extinction of garlic mustard (Alliaria petiolata) populations: implications for weed science and conservation biology

Abstract: An assumption of weed science and conservation biology is that small populations are more vulnerable to elimination and extinction than large populations We tested this with the invasive biennial garlic mustard (Alliaria petiolata) We compared 61 experimental populations from which every flowering plant was removed for 4 years, with 56 control populations Whereas the majority of the control populations continued to expand in size over the 4 years, experimental populations showed a strong experimental effect, remaining stable in size, declining in size, or going extinct Small populations were far more vulnerable to extinction than large populations: 43% of small experimental populations (initially fewer than 10 individuals) went extinct, but only 7% of large populations (initially more than 50 individuals) However, some small experimental populations persisted, and in a few cases, larger experimental populations continued to expand even though every flowering individual had been removed These results and a simple population model suggest the importance of buried seeds in allowing this species to persist despite attempts to eradicate it

The assembly of an island fauna by natural invasion: Sources and temporal patterns in the avian colonization of Barbados

Abstract: By virtue of their isolation and depauperate faunas, oceanic islands offer unique opportunities to characterize the historical development of ecological communities derived from both natural and anthropogenic invasions. Barbados, an outlying island in the Lesser Antilles, was formed approximately 700,000 YBP by tectonic uplift and was then colonized by birds via natural invasion from the much older volcanic islands in the main Lesser Antillean arc. We investigated the timing and sources of the avian invasion of Barbados by determining levels of mitochondrial DNA (mtDNA) divergence between populations of eight bird species from Barbados and those on the nearby putative source islands of St. Lucia and St. Vincent. Although all Barbados populations appeared to be young relative to the geological age of the island, we found differences among species in their inferred times of colonization and we identified at least two sources of immigrants to Barbados. In contrast to these historical differences across species and populations, our characterization of the mitochondrial genotypes of 231 individual birds suggests that each island population represents the descendants of a single founding maternal lineage. Considered in concert, the results of this molecular survey indicate that the Barbados bird community is composed of species with different invasion histories, which in turn suggests that the island’s community composition has changed repeatedly over its 700,000 year history.

The Global Invasive Species Program (GISP)

Abstract: During the period between 1982 and 1988 the Scientific Committee on Problems of the Environment (SCOPE) engaged a large number of scientists in an effort to document the nature of the invasive species ‘problem’. The results of this effort appeared in a number of books and as a synthesis in 1989 entitled Biological Invasions: A Global Perspective (Drake et al., 1989). This synthesis clearly established that invasive species could have major impacts on ecosystem functioning and that virtually all ecosystems were impacted by them, even those under preservation management. It was also clear that we are on the way to establishing a whole new biotic order on the Earth due to the massive breakdown of biogeographic barriers to migration. Although the SCOPE programme was quite successful scientifically it did not offer much to managers except to inform them they were not alone in the world dealing with these problems. SCOPE, along with partners from UNEP (United Nations Environment Program), IUCN (International Union for the Conservation of Nature) and CABI (Commonwealth Agricultural Bureau International), is embarking on a new programme on invasive species, this time with the explicit objective of providing new tools for understanding as well as dealing with invasive species. This venture is called the Global Invasive Species Program, or GISP. This effort differs substantially from the previous programme in that it will engage the many constituencies that are involved with the problem including natural and social scientists, educators, lawyers, resource managers, and people from both industry and government. The programme has 11 elements. Each of these will contribute to building the comprehensive approach that is needed for dealing with invasive species. Four elements deal with synthesizing our current knowledge on invasive species. These include the ecology of invasive species (led by David Richardson

Genetic and Environmental Controls on the Establishment of the Invasive Grass, Phalaris arundinacea

Abstract: Whether an exotic species becomes integrated into a community or aggressively takes it over depends upon many interacting factors. Using contextual analyses, we combined genetic data about an invasive plant with information about the neighboring species, the community, and the environment to determine what factors enable a genotype or species to invade. We transplanted 50 individuals of each of three clones of the invasive grass Phalaris arundinacea, reed canary grass, into 150 random locations within a Vermont pasture. For each individual, we recorded clonal identity, neighbor identity, community indices (species richness and species diversity), and an environmental variable (soil moisture). The response variables were survivorship, above-ground biomass, below-ground biomass and the ratio of above- to below-ground biomass. Clonal identity affected both survivorship and below-ground biomass. The fastest tillering clone had poor survivorship but survivors produced a large amount of below-ground biomass, making this clone more likely to successfully overwinter. Neighbor species affected above- and below-ground biomass. Reed canary grass produced more above- and below-ground biomass when Anthoxanthum odoratum, a common pasture grass species, was abundant. Community attributes also influenced growth. Although we expected diverse plots to repel the invasion, plants in the more diverse plots had higher amounts of below-ground biomass. Finally, environmental effects also influenced growth. Reed canary grass produced more above-ground biomass in wetter plots, confirming that it does well under wet conditions.