Minimizing opportunity costs to aquatic connectivity restoration while controlling an invasive species

Controlling invasive species is critical for conservation but can have unintended consequences for native species and divert resources away from other efforts. This dilemma occurs on a grand scale in the North American Great Lakes, where dams and culverts block tributary access to habitat of desirable fish species and are a lynchpin of long‐standing efforts to limit ecological damage inflicted by the invasive, parasitic sea lamprey (Petromyzon marinus). Habitat restoration and sea‐lamprey control create conflicting goals for managing aging infrastructure. We used optimization to minimize opportunity costs of habitat gains for 37 desirable migratory fishes that arose from restricting sea lamprey access (0–25% increase) when selecting barriers for removal under a limited budget (US$1–105 million). Imposing limits on sea lamprey habitat reduced gains in tributary access for desirable species by 15–50% relative to an unconstrained scenario. Additional investment to offset the effect of limiting sea‐lamprey access resulted in high opportunity costs for 30 of 37 species (e.g., an additional US$20–80 million for lake sturgeon [Acipenser fulvescens]) and often required ≥5% increase in sea‐lamprey access to identify barrier‐removal solutions adhering to the budget and limiting access. Narrowly distributed species exhibited the highest opportunity costs but benefited more at less cost when small increases in sea‐lamprey access were allowed. Our results illustrate the value of optimization in limiting opportunity costs when balancing invasion control against restoration benefits for diverse desirable species. Such trade‐off analyses are essential to the restoration of connectivity within fragmented rivers without unleashing invaders.     

The use of environmental DNA in invasive species surveillance of the Great Lakes commercial bait trade

Over 180 non‐native species have been introduced in the Laurentian Great Lakes region, many posing threats to native species and ecosystem functioning. One potential pathway for introductions is the commercial bait trade; unknowing or unconcerned anglers commonly release unused bait into aquatic systems. Previous surveillance efforts of this pathway relied on visual inspection of bait stocks in retail shops, which can be time and cost prohibitive and requires a trained individual that can rapidly and accurately identify cryptic species. Environmental DNA (eDNA) surveillance, a molecular tool that has been used for surveillance in aquatic environments, can be used to efficiently detect species at low abundances. We collected and analyzed 576 eDNA samples from 525 retail bait shops throughout the Laurentian Great Lake states. We used eDNA techniques to screen samples for multiple aquatic invasive species (AIS) that could be transported in the bait trade, including bighead (Hypophthalmichthys nobilis) and silver carp (H. molitrix), round goby (Neogobius melanostomus), tubenose goby (Proterorhinus marmoratus), Eurasian rudd (Scardinius erythrophthalmus), and goldfish (Carassius auratus). Twenty‐seven samples were positive for at least one target species (4.7% of samples), and all target species were found at least once, except bighead carp. Despite current regulations, the bait trade remains a potential pathway for invasive species introductions in the Great Lakes region. Alterations to existing management strategies regarding the collection, transportation, and use of live bait are warranted, including new and updated regulations, to prevent future introductions of invasive species in the Great Lakes via the bait trade. El Uso del ADN Ambiental en la Vigilancia de Especies Invasoras del Mercado de Carnada Comercial de los Grandes Lagos     

Optimisation of adjusted barrier management to improve glass eel migration at an estuarine barrier

Estuarine barriers may significantly reduce the upstream migration of diadromous fish species like the European eel (Anguilla anguilla L.). Previous research showed that limited barrier opening during tidal rise was a cost-efficient and effective mitigation option to improve upstream glass eel migration, without significant intrusion of sea water. This paper aims to optimise this adjusted barrier management to improve eel passage at a tidal barrier complex at the mouth of the River Yser, Flanders, Belgium, one of the most important migration routes for glass eel in Flanders. Specifically, three hypotheses were tested. The first hypothesis analysed the impact of the number of barriers opened on the upstream glass eel migration. The second hypothesis evaluated the relation between the size of the barrier opening and glass eel migration. Finally, we tested whether the suggested adjusted barrier management may lead to a significant increase in conductivity in the River Yser. Increased opening of one barrier appeared more efficient than opening several barriers slightly. Conductivity increased during periods of extreme drought and at base flow, but decreased within 24 h after the first peak flow. This indicates that adjusted barrier management does not entail salt intrusion in the Yser basin, as long as this management is not applied in extremely dry periods. Since the adjusted barrier management is easily implemented and could be applied on numerous tidal barriers, the presented results may contribute to restoration of eel populations worldwide and be of interest to a wide range of river managers and stakeholders.     

Fecundity as a Basis for Risk Assessment of Nonindigenous Freshwater Molluscs

Abstract:  The most efficient way to reduce future damages from nonindigenous species is to prevent the introduction of harmful species. Although ecologists have long sought to predict the identity of such species, recent methodological advances promise success where previous attempts failed. We applied recently developed risk assessment approaches to nonindigenous freshwater molluscs at two geographic scales: the Laurentian Great Lakes basin and the 48 contiguous states of the United States. We used data on natural history and biogeography to discriminate between established freshwater molluscs that are benign and those that constitute nuisances (i.e., cause environmental and/or economic damage). Two statistical techniques, logistic regression and categorical tree analysis, showed that nuisance status was positively associated with fecundity. Other aspects of natural history and biogeography did not significantly affect likelihood of becoming a nuisance. We then used the derived statistical models to predict the chance that 15 mollusc species not yet in natural ecosystems would cause damage if they become established. We also tested whether time since establishment is related to the likelihood that nonindigenous mollusc species in the Great Lakes and United States would cause negative impacts. No significant relationship was evident at the U.S. scale, but recently established molluscs within the Great Lakes were more likely to cause negative impacts. This may reflect changing environmental conditions, changing patterns of trade, or may be an indication of "invasional meltdown." Our quantitative analyses could be extended to other taxa and ecosystems and offer a number of improvements over the qualitative risk assessments currently used by U.S. (and other) government agencies.     

Past,present, and future of a freshwater fish metapopulation in a threatened landscape

It is well documented that hydropower plants can affect the dynamics of fish populations through landscape alterations and the creation of new barriers. Less emphasis has been placed on the examination of the genetic consequences for fish populations of the construction of dams. The relatively few studies that focus on genetics often do not consider colonization history and even fewer tend to use this information for conservation purposes. As a case study, we used a 3‐pronged approach to study the influence of historical processes, contemporary landscape features, and potential future anthropogenic changes in landscape on the genetic diversity of a fish metapopulation. Our goal was to identify the metapopulation's main attributes, detect priority areas for conservation, and assess the consequences of the construction of hydropower plants for the persistence of the metapopulation. We used microsatellite markers and coalescent approaches to examine historical colonization processes, traditional population genetics, and simulations of future populations under alternate scenarios of population size reduction and gene flow. Historical gene flow appeared to have declined relatively recently and contemporary populations appeared highly susceptible to changes in landscape. Gene flow is critical for population persistence. We found that hydropower plants could lead to a rapid reduction in number of alleles and to population extirpation 50–80 years after their construction. More generally, our 3‐pronged approach for the analyses of empirical genetic data can provide policy makers with information on the potential impacts of landscape changes and thus lead to more robust conservation efforts.     

Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change

Ecosystem change often affects the structure of aquatic communities thereby regulating how much and by what pathways energy and critical nutrients flow through food webs. The availability of energy and essential nutrients to top predators such as seabirds that rely on resources near the water's surface will be affected by changes in pelagic prey abundance. Here, we present results from analysis of a 25-year data set documenting dietary change in a predatory seabird from the Laurentian Great Lakes. We reveal significant declines in trophic position and alterations in energy and nutrient flow over time. Temporal changes in seabird diet tracked decreases in pelagic prey fish abundance. As pelagic prey abundance declined, birds consumed less aquatic prey and more terrestrial food. This pattern was consistent across all five large lake ecosystems. Declines in prey fish abundance may have primarily been the result of predation by stocked piscivorous fishes, but other lake-specific factors were likely also important. Natural resource management activities can have unintended consequences for nontarget ecosystem components. Reductions in pelagic prey abundance have reduced the capacity of the Great Lakes to support the energetic requirements of surface-feeding seabirds. In an environment characterized by increasingly limited pelagic fish resources, they are being offered a Hobsonian choice: switch to less nutritious terrestrial prey or go hungry.     

Evaluating complementary networks of restoration plantings for landscape‐scale occurrence of temporally dynamic species

Multibillion dollar investments in land restoration make it critical that conservation goals are achieved cost‐effectively. Approaches developed for systematic conservation planning offer opportunities to evaluate landscape‐scale, temporally dynamic biodiversity outcomes from restoration and improve on traditional approaches that focus on the most species‐rich plantings. We investigated whether it is possible to apply a complementarity‐based approach to evaluate the extent to which an existing network of restoration plantings meets representation targets. Using a case study of woodland birds of conservation concern in southeastern Australia, we compared complementarity‐based selections of plantings based on temporally dynamic species occurrences with selections based on static species occurrences and selections based on ranking plantings by species richness. The dynamic complementarity approach, which incorporated species occurrences over 5 years, resulted in higher species occurrences and proportion of targets met compared with the static complementarity approach, in which species occurrences were taken at a single point in time. For equivalent cost, the dynamic complementarity approach also always resulted in higher average minimum percent occurrence of species maintained through time and a higher proportion of the bird community meeting representation targets compared with the species‐richness approach. Plantings selected under the complementarity approaches represented the full range of planting attributes, whereas those selected under the species‐richness approach were larger in size. Our results suggest that future restoration policy should not attempt to achieve all conservation goals within individual plantings, but should instead capitalize on restoration opportunities as they arise to achieve collective value of multiple plantings across the landscape. Networks of restoration plantings with complementary attributes of age, size, vegetation structure, and landscape context lead to considerably better outcomes than conventional restoration objectives of site‐scale species richness and are crucial for allocating restoration investment wisely to reach desired conservation goals.     

Indirect upstream effects of dams: consequences of migratory consumer extirpation in Puerto Rico.

Large dams degrade the integrity of a wide variety of ecosystems, yet direct downstream effects of dams have received the most attention from ecosystem managers and researchers. We investigated indirect upstream effects of dams resulting from decimation of migratory freshwater shrimp and fish populations in Puerto Rico, USA, in both high- and low-gradient streams. In high-gradient streams above large dams, native shrimps and fishes were extremely rare, whereas similar sites without large dams had high abundances of native consumers. Losses of native fauna above dams dramatically altered their basal food resources and assemblages of invertebrate competitors and prey. Compared to pools in high-gradient streams with no large dams, pool epilithon above dams had nine times more algal biomass, 20 times more fine benthic organic matter (FBOM), 65 times more fine benthic inorganic matter (FBIM), 28 times more carbon, 19 times more nitrogen, and four times more non-decapod invertebrate biomass. High-gradient riffles upstream from large dams had five times more FBIM than did undammed riffles but showed no difference in algal abundance, FBOM, or non-decapod invertebrate biomass. For epilithon of low-gradient streams, differences in basal resources between pools above large dams vs. without large dams were considerably smaller in magnitude than those observed for pools in high-gradient sites. These results match previous stream experiments in which the strength of native shrimp and fish effects increased with stream gradient. Our results demonstrate that dams can indirectly affect upstream free-flowing reaches by eliminating strong top-down effects of consumers. Migratory omnivorous shrimps and fishes occur throughout the tropics, and the consequences of their declines upstream from many tropical dams are likely to be similar to those in Puerto Rico. Thus, ecological effects of migratory fauna loss upstream from dams encompass a wider variety of species interactions and biomes than the bottom-up effects (i.e., elimination of salmonid nutrient subsidies) recognized for northern temperate systems.     

Predicting barrier passage and habitat suitability for migratory fish species

Fish migrate to spawn, feed, seek refuge from predators, and escape harmful environmental conditions. The success of upstream migration is limited by the presence of barriers that can impede the passage of fish. We used a spatially explicit modeling strategy to examine the effects of barriers on passage for 21 native and non-native migratory fish species and the amount of suitable habitat blocked for each species. Spatially derived physical parameter estimates and literature based fish capabilities and tolerances were used to predict fish passage success and habitat suitability. Both the fish passage and the habitat suitability models accurately predicted fish presence above barriers for most common, non-stocked species. The fish passage model predicted that barriers greater than or equal to 6 m block all migratory species. Chinook salmon (Oncorhynchus tshawytscha) was expected to be blocked the least. The habitat suitability model predicted that low gradient streams with intact habitat quality were likely to support the highest number of fish species. The fish passage and habitat suitability models were intended to be used by environmental managers as strategy development tools to prioritize candidate dams for field assessment and make decisions regarding the management of migratory fish populations.     

Discard analysis and damage to non-target species in the "rapido" trawl fishery

This paper describes the catch composition in the rapido trawl fishery and the direct effects on non-target species. All data were collected on commercial fishing vessels so as to reflect commercial rapido-trawling practice. The effects on non-target species were measured using two different damage scales (three- and seven-level scales) depending on the morphology of the taxa. Damage assessment was performed taking into account the whole fishing process by collecting individuals that passed through the cod-end, individuals that were retained in the cod-end and dropped onto the deck and individuals that were collected at the end of the sorting operation just before their return to the sea. Due to differences in the habitat and spatial distribution of target species, discard/commercial ratio was very different among the three different target species fisheries: 1:6 in the queen scallop (Aequipecten opercularis) fishery, 2:1 in the flatfish (Solea spp., Platichthys flesus, Psetta maximus and Scophthalmus rhombus) fishery and 9:1 in the scallop (Pecten jacobaeus) fishery. Damage sustained by non-target species was species-specific and related to the morphology of different organisms. The sorting operation produced similar levels of injury to those of the gear itself: all discarded animals showed higher levels of damage after the sorting than before. Damage to animals that had passed through the cod-end followed the same pattern, and these data could give an estimate of the "unobserved mortality". Our observations indicated a higher impact on non-target species caused by the queen scallop fishery than that caused by the flatfish fishery. This could be due to the total amount of hard-shelled species (in the queen scallop fishery, A. opercularis accounted for 87% of the total catch biomass) in any given haul, since shells macerated the catch during towing. Discarded animals from the queen scallop fishery showed higher levels of damage than those collected in the flatfish fishery. The rapido trawl fishery seemed to exert a strong selective pressure on the macrobenthic community, being able to modify the epibenthic fauna structure which, in heavily exploited fishing grounds, was dominated by bivalves, gastropods, crabs, starfish and brittlestars.     

Effects of urbanization and urban stream restoration on the physical and biological structure of stream ecosystems

Streams, as low-lying points in the landscape, are strongly influenced by the stormwaters, pollutants, and warming that characterize catchment urbanization. River restoration projects are an increasingly popular method for mitigating urban insults. Despite the growing frequency and high expense of urban stream restoration projects, very few projects have been evaluated to determine whether they can successfully enhance habitat structure or support the stream biota characteristic of reference sites. We compared the physical and biological structure of four urban degraded, four urban restored, and four forested streams in the Piedmont region of North Carolina to quantify the ability of reach-scale stream restoration to restore physical and biological structure to urban streams and to examine the assumption that providing habitat is sufficient for biological recovery. To be successful at mitigating urban impacts, the habitat structure and biological communities found in restored streams should be more similar to forested reference sites than to their urban degraded counterparts. For every measured reach- and patch-scale attribute, we found that restored streams were indistinguishable from their degraded urban stream counterparts. Forested streams were shallower, had greater habitat complexity and median sediment size, and contained less-tolerant communities with higher sensitive taxa richness than streams in either urban category. Because heavy machinery is used to regrade and reconfigure restored channels, restored streams had less canopy cover than either forested or urban streams. Channel habitat complexity and watershed impervious surface cover (ISC) were the best predictors of sensitive taxa richness and biotic index at the reach and catchment scale, respectively. Macroinvertebrate communities in restored channels were compositionally similar to the communities in urban degraded channels, and both were dissimilar to communities in forested streams. The macroinvertebrate communities of both restored and urban degraded streams were correlated with environmental variables characteristic of degraded urban systems. Our study suggests that reach-scale restoration is not successfully mitigating for the factors causing physical and biological degradation.     

Boundaries and Corridors as a Continuum of Ecological Flow Control: Lessons from Rivers and Streams

Abstract: Landscape boundaries and corridors are areas of small spatial extent relative to their large effects on ecological flows. The trend in ecological literature is to treat corridors and boundaries as separate phenomena on the landscape. This approach, however, misses a fundamental aspect they have in common: their strong influence on ecological flows. Corridors and boundaries exist at opposite ends of a permeability gradient, differing in their effects on rates and direction of flow. The position of landscape structures along this permeability gradient depends on attributes of both the flow and of the structure itself. We discuss boundaries and corridors in terms of mover specificity, scale, and effects on different levels of ecological organization, using rivers and streams to illustrate our points. We predict which structures will act as boundaries or corridors and at what spatial and temporal scales they are likely to be relevant. Considering the function of landscape structures across the boundary-corridor continuum will provide researchers and managers with a more complete, holistic viewpoint and will allow better strategies to attain conservation goals.     

Fish dispersal in fragmented landscapes: a modeling framework for quantifying the permeability of structural barriers

Dispersal is a key determinant of the spatial distribution and abundance of populations, but human-made fragmentation can create barriers that hinder dispersal and reduce population viability. This study presents a modeling framework based on dispersal kernels (modified Laplace distributions) that describe stream fish dispersal in the presence of obstacles to passage. We used mark-recapture trials to quantify summer dispersal of brook trout (Salvelinus fontinalis) in four streams crossed by a highway. The analysis identified population heterogeneity in dispersal behavior, as revealed by the presence of a dominant sedentary component (48-72% of all individuals) characterized by short mean dispersal distance (<10 m), and a secondary mobile component characterized by longer mean dispersal distance (56-1086 m). We did not detect evidence of barrier effects on dispersal through highway crossings. Simulation of various plausible scenarios indicated that detectability of barrier effects was strongly dependent on features of sampling design, such as spatial configuration of the sampling area, barrier extent, and sample size. The proposed modeling framework extends conventional dispersal kernels by incorporating structural barriers. A major strength of the approach is that ecological process (dispersal model) and sampling design (observation model) are incorporated simultaneously into the analysis. This feature can facilitate the use of prior knowledge to improve sampling efficiency of mark-recapture trials in movement studies. Model-based estimation of barrier permeability and its associated uncertainty provides a rigorous approach for quantifying the effect of barriers on stream fish dispersal and assessing population dynamics of stream fish in fragmented landscapes.     

Invertebrate community responses to recreational clam digging

Marine reserves can help in maintaining biodiversity and potentially be useful as a fishery management tool by removing human-mediated impacts. Intertidal, soft-sediment habitats can often support robust recreational and commercial shellfish harvests, especially for clams; however, there is limited research on the effects of reserves in these habitats. In San Juan County, Washington, several reserves prohibit recreational clam digging. We examined the effects of these reserves on infaunal community composition through comparison with non-reserve beaches during a 6-week period. Clam abundance, overall species richness and total polychaete family richness were greater on reserve beaches compared to non-reserve beaches. Additionally, an experiment within a reserve demonstrated negative impacts of digging on non-target infauna. These effects probably resulted from local disruption and disturbance of the sediment habitat and not from increased post-digging predation, which was controlled. Intertidal reserves could play an important role in sustaining local and potentially regional biodiversity.     

Characteristics of Important Stopover Locations for Migrating Birds: Remote Sensing with Radar in the Great Lakes Basin

Abstract:  A preliminary stage in developing comprehensive conservation plans involves identifying areas used by the organisms of interest. The areas used by migratory land birds during temporal breaks in migration (stopover periods) have received relatively little research and conservation attention. Methodologies for identifying stopover sites across large geographic areas have been, until recently, unavailable. Advances in weather-radar technology now allow for evaluation of bird migration patterns at large spatial scales. We analyzed radar data (WSR-88D) recorded during spring migration in 2000 and 2001 at 6 sites in the Great Lakes basin (U.S.A.). Our goal was to link areas of high migrant activity with the land-cover types and landscape contexts corresponding to those areas. To characterize the landscapes surrounding stopover locations, we integrated radar and land-cover data within a geographic information system. We compared landscape metrics within 5 km of areas that consistently hosted large numbers of migrants with landscapes surrounding randomly selected areas that were used by relatively few birds during migration. Concentration areas were characterized by 1.2 times more forest cover and 9.3 times more water cover than areas with little migrant activity. We detected a strong negative relationship between activity of migratory birds and agricultural land uses. Examination of individual migration events confirmed the importance of fragments of forested habitat in highly altered landscapes and highlighted large concentrations of birds departing from near-shore terrestrial areas in the Great Lakes basin. We conclude that conservation efforts can be more effectively targeted through intensive analysis of radar imagery.     

Recruitment of Hexagenia mayfly nymphs in western Lake Erie linked to environmental variability.

After a 40-year absence caused by pollution and eutrophication, burrowing mayflies (Hexagenia spp.) recolonized western Lake Erie in the mid 1990s as water quality improved. Mayflies are an important food resource for the economically valuable yellow perch fishery and are considered to be major indicator species of the ecological condition of the lake. Since their reappearance, however, mayfly populations have suffered occasional unexplained recruitment failures. In 2002, a failure of fall recruitment followed an unusually warm summer in which western Lake Erie became temporarily stratified, resulting in low dissolved oxygen levels near the lake floor. In the present study, we examined a possible link between Hexagenia recruitment and periods of intermittent stratification for the years 1997 2002. A simple model was developed using surface temperature, wind speed, and water column data from 2003 to predict stratification. The model was then used to detect episodes of stratification in past years for which water column data are unavailable. Low or undetectable mayfly recruitment occurred in 1997 and 2002, years in which there was frequent or extended stratification between June and September. Highest mayfly reproduction in 2000 corresponded to the fewest stratified periods. These results suggest that even relatively brief periods of stratification can result in loss of larval mayfly recruitment, probably through the effects of hypoxia. A trend toward increasing frequency of hot summers in the Great Lakes region could result in recurrent loss of mayfly larvae in western Lake Erie and other shallow areas in the Great Lakes.     

Effects of Dam‐Induced Landscape Fragmentation on Amazonian Ant–Plant Mutualistic Networks

Mutualistic networks are critical to biological diversity maintenance; however, their structures and functionality may be threatened by a swiftly changing world. In the Amazon, the increasing number of dams poses a large threat to biological diversity because they greatly alter and fragment the surrounding landscape. Tight coevolutionary interactions typical of tropical forests, such as the ant–myrmecophyte mutualism, where the myrmecophyte plants provide domatia nesting space to their symbiotic ants, may be jeopardized by the landscape changes caused by dams. We analyzed 31 ant–myrmecophyte mutualistic networks in undisturbed and disturbed sites surrounding Balbina, the largest Central Amazonian dam. We tested how ant–myrmecophyte networks differ among dam‐induced islands, lake edges, and undisturbed forests in terms of species richness, composition, structure, and robustness (number of species remaining in the network after partner extinctions). We also tested how landscape configuration in terms of area, isolation, shape, and neighborhood alters the structure of the ant–myrmecophyte networks on islands. Ant–myrmecophytic networks were highly compartmentalized in undisturbed forests, and the compartments had few strongly connected mutualistic partners. In contrast, networks at lake edges and on islands were not compartmentalized and were negatively affected by island area and isolation in terms of species richness, density, and composition. Habitat loss and fragmentation led to coextinction cascades that contributed to the elimination of entire ant–plant compartments. Furthermore, many myrmecophytic plants in disturbed sites lost their mutualistic ant partners or were colonized by opportunistic, nonspecialized ants. Robustness of ant–myrmecophyte networks on islands was lower than robustness near lake edges and in undisturbed forest and was particularly susceptible to the extinction of plants. Beyond the immediate habitat loss caused by the building of large dams in Amazonia, persistent edge effects and habitat fragmentation associated with dams had large negative effects on animal–plant mutualistic networks. Efectos de la Fragmentación del Paisaje Inducida por Presas sobre Redes Mutualistas Hormiga‐Planta Amazónicas     

Predicting invasion risk using measures of introduction effort and environmental niche models.

The Chinese mitten crab (Eriocheir sinensis) is native to east Asia, is established throughout Europe, and is introduced but geographically restricted in North America. We developed and compared two separate environmental niche models using genetic algorithm for rule set prediction (GARP) and mitten crab occurrences in Asia and Europe to predict the species' potential distribution in North America. Since mitten crabs must reproduce in water with >15% per hundred salinity, we limited the potential North American range to freshwater habitats within the highest documented dispersal distance (1260 km) and a more restricted dispersal limit (354 km) from the sea. Applying the higher dispersal distance, both models predicted the lower Great Lakes, most of the eastern seaboard, the Gulf of Mexico and southern extent of the Mississippi River watershed, and the Pacific northwest as suitable environment for mitten crabs, but environmental match for southern states (below 35 degrees N) was much lower for the European model. Use of the lower range with both models reduced the expected range, especially in the Great Lakes, Mississippi drainage, and inland areas of the Pacific Northwest. To estimate the risk of introduction of mitten crabs, the amount of reported ballast water discharge into major United States ports from regions in Asia and Europe with established mitten crab populations was used as an index of introduction effort. Relative risk of invasion was estimated based on a combination of environmental match and volume of unexchanged ballast water received (July 1999-December 2003) for major ports. The ports of Norfolk and Baltimore were most vulnerable to invasion and establishment, making Chesapeake Bay the most likely location to be invaded by mitten crabs in the United States. The next highest risk was predicted for Portland, Oregon. Interestingly, the port of Los Angeles/Long Beach, which has a large shipping volume, had a low risk of invasion. Ports such as Jacksonville, Florida, had a medium risk owing to small shipping volume but high environmental match. This study illustrates that the combination of environmental niche- and vector-based models can provide managers with more precise estimates of invasion risk than can either of these approaches alone.     

Spatial heterogeneity influences native and nonnative plant species richness

Spatial heterogeneity may have differential effects on the distribution of native and nonnative plant species richness. We examined the effects of spatial heterogeneity on native and nonnative plant species richness distributions in the central part of Rocky Mountain National Park, Colorado, USA. Spatial heterogeneity around vegetation plots was characterized using landscape metrics, environmental/topographic variables (slope, aspect, elevation, and distance from stream or river), and soil variables (nitrogen, clay, and sand). The landscape metrics represented five components of landscape heterogeneity and were measured at four spatial extents (within varying radii of 120, 240, 480, and 960 m) using the FRAGSTATS landscape pattern analysis program. Akaike's Information Criterion adjusted for small sample size (AICc) was used to select the best models from a set of multiple linear regression models developed for native and nonnative plant species richness at four spatial extents and three levels of ecological hierarchy (i.e., landscape, land cover, and community). Both native and nonnative plant species richness were positively correlated with edge density, Simpson's diversity index and interspersion/juxtaposition index, and were negatively correlated with mean patch size. The amount of variation explained at four spatial extents and three hierarchical levels ranged from 30% to 70%. At the landscape level, the best models explained 43% of the variation in native plant species richness and 70% of the variation in nonnative plant species richness (240-m extent). In general, the amount of variation explained was always higher for nonnative plant species richness, and the inclusion of landscape metrics always significantly improved the models. The best models explained 66% of the variation in nonnative plant species richness for both the conifer land cover type and lodgepole pine community. The relative influence of the components of spatial heterogeneity differed for native and nonnative plant species richness and varied with the spatial extent of analysis and levels of ecological hierarchy. The study offers an approach to quantify spatial heterogeneity to improve models of plant biodiversity. The results demonstrate that ecologists must recognize the importance of spatial heterogeneity in managing native and nonnative plant species.