Pathways and places associated with nonindigenous aquatic species introductions in the Laurentian Great Lakes

The present study aimed to evaluate the health status of Octopus maya females on different days after spawning (days 0, 10, 20, 30, and 40). A total of 25 O. maya females were examined in terms of physiological (i.e., weight loss, hepatosomatic and gonadosomatic indexes, and hemocyanin, protein, glucose, cholesterol, and acylglycerides concentrations in plasma) and immunological variables (i.e., total hemocyte count, hemagglutination, and phenoloxidase activity). We hypothesized that O. maya females should maintain their physiological integrity throughout the post-spawning period until the hatching of the offspring. Results showed that the physiological and immunological indicators measured in post-spawning females significantly changed with time. Loss of body weight over time and a decrease in the hepatosomatic and gonadosomatic indexes were observed. Hemolymph components showed variations that reflect the consumption of reserves and coincide with an increased immune process of hemagglutination and phenoloxidase activity in hemocytes. Our results demonstrate that O. maya females are adapted to maintain an adequate state of health to care for their spawn despite the long period of starvation and contribute to the identification of the mechanisms involved in maintaining the integrity of these animals during one of the most critical phases of their life cycle.     

Environmental DNA detection of rare and invasive fish species in two Great Lakes tributaries

The extraction and characterization of DNA from aquatic environmental samples offers an alternative, noninvasive approach for the detection of rare species. Environmental DNA, coupled with PCR and next‐generation sequencing (“metabarcoding”), has proven to be very sensitive for the detection of rare aquatic species. Our study used a custom‐designed group‐specific primer set and next‐generation sequencing for the detection of three species at risk (Eastern Sand Darter, Ammocrypta pellucida; Northern Madtom, Noturus stigmosus; and Silver Shiner, Notropis photogenis), one invasive species (Round Goby, Neogobius melanostomus) and an additional 78 native species from two large Great Lakes tributary rivers in southern Ontario, Canada: the Grand River and the Sydenham River. Of 82 fish species detected in both rivers using capture‐based and eDNA methods, our eDNA method detected 86.2% and 72.0% of the fish species in the Grand River and the Sydenham River, respectively, which included our four target species. Our analyses also identified significant positive and negative species co‐occurrence patterns between our target species and other identified species. Our results demonstrate that eDNA metabarcoding that targets the fish community as well as individual species of interest provides a better understanding of factors affecting the target species spatial distribution in an ecosystem than possible with only target species data. Additionally, eDNA is easily implemented as an initial survey tool, or alongside capture‐based methods, for improved mapping of species distribution patterns.     

Towards defining aquatic ecosystem health for the Great Lakes

The Canada — U.S. Great Lakes Water Quality Agreement defines Areas of Concern as geographic areas that fail to meet the general or specific objectives of the Great Lakes Water Quality Agreement where such failure has caused or is likely to cause impairment of beneficial use or the area's ability to support aquatic life. Impairment of beneficial use is defined by the Agreement as a change in the physical, chemical or biological integrity sufficient to cause any one of 14 designated use impairments. In 1987 the International Joint Commission's Great Lakes Water Quality Board (GLWQB) recommended that criteria be developed to determine when ecosystem conditions have been impacted enough to warrant designation as an Area of Concern and when conditions have improved sufficiently to be delisted. Based on scientific input and policy considerations, the GLWQB adopted, in principle, a set of quantitative and qualitative listing/delisting criteria for each of the 14 use impairments. These criteria can be uniformly applied throughout the basin. Further, the GLWQB recommended future refinement of these criteria based on advances in science and public input.     

Wetland food resources for spring-migrating ducks in the Upper Mississippi River and Great Lakes Region

Wetlands in the Upper Mississippi River and Great Lakes Region (UMRGLR) must annually sustain populations of migrating waterfowl from the mid-continent of North America. We used multi-stage sampling to estimate plant and invertebrate food biomasses (kg/ha) for ducks in 3 wetland habitat types at 6 stop-over locations in the UMRGLR during 2006 and 2007. Total biomass was greatest in palustrine emergent (PEM; = 208 kg/ha, SE = 23, median = 120), followed by palustrine forested (PF; = 87 kg/ha, SE = 7; median = 43), and lacustrine–riverine (LR; = 52 kg/ha, SE = 7; median = 27) wetlands. Ducks that foraged in forested and LR wetlands encountered the least food abundance during spring in the UMRGLR. Our estimates of food abundance were the lowest reported among other landscape scale surveys from mid-continent North America. About 1 in every 5 PEM wetlands and over half of our PF and LR wetlands that we sampled contained <50 kg/ha of food, suggesting many had little or no forage value to ducks during spring. Biomass of plant foods generally exceeded invertebrate biomass in all habitat types, although invertebrate biomass estimates exceeded plant biomass in 8 of 29 sites when considered by wetland type and year. Total food biomass estimates varied widely ( = 6–425 kg/ha) between years and among habitats; thus, using global arithmetic means to estimate food abundance for conservation planning obscures fine scale temporal and spatial variation that may be necessary for management on local and sub-regional levels. Distributions of food biomass estimates were right-skewed, causing us to question whether arithmetic means realistically represent levels of food abundance that all ducks encounter during spring migration. Alternative measures of central tendency (e.g., median) may be more biologically realistic, particularly if spring-migrating ducks are not distributed in an ideal-free manner with respect to food abundance. Future research should determine how ducks distribute themselves in relation to variation in food abundance in space and time during spring migration to strengthen the biological approach to conservation planning in non-breeding Joint Venture areas of the North American Waterfowl Management Plan. © 2011 The Wildlife Society.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Evaluating management options for aquatic invasive species: concepts and methods

In analyses that attempt to estimate the costs of species invasions, it has been typical to report the costs of management and/or to multiply per-unit costs by the number affected to arrive at a total. These estimates are of limited value for most policy questions. We start our discussion by recognizing that biological pollutants such as aquatic invasive species are like conventional pollutants in important ways and appeal to the well-developed literature on conventional pollution to guide our thinking into how best to conceptualize the problem. We use a standard pollution control framework to identify the margins over which costs and benefits should be estimated to guide wise decision-making. We then use examples from the literature to illustrate how transactions in related markets can be used to estimate the benefits of management. The roles of adaptation, mitigation, and species population growth have particular relevance and are highlighted. In the final section of the paper, we think through the conditions under which investing in genetic biocontrol methods would be economically justified.     

Vulnerability of benthic habitats to the aquatic invasive species

A comparative vulnerability analysis of 16 selected benthic habitat types in the SE Baltic Sea waters and the Curonian lagoon, including Klaipeda strait, was performed using long-term monitoring datasets (1980–2003) and results of several other surveys in the lagoon and the sea. Results indicated that invasive species richness (number of alien species per habitat) in lagoon habitats was significantly higher than in the sea. Habitats formed by artificial rock and stone, sand, mud, and habitats modified by zebra mussel shell deposits appeared to be the most invaded. Highest invasive species richness occurred in habitats with high native species richness indicating that the main factors driving native species distribution (such as favourable physical conditions, habitat alterations generated by human or/and biotic activities) are also driving aquatic invaders. Physical factors distinguished to be the most important for native and invasive species distribution were salinity, depth range (expressed by the maximal and minimal depths difference within a habitat), shallowness of a habitat (expressed by a minimal depth), and availability of a hard substrate.     

Estimating the financial costs of freshwater invasive species in Great Britain: a standardized approach to invasive species costing

Both ecological and economic impacts factor into invasive alien species (IAS) management considerations; however, economic impacts are often difficult to assess, much less quantify. Studies frequently aggregate identified financial costs as a proxy for IAS economic impacts, but these aggregate figures are often generated in an ad hoc fashion. Such estimates typically sum disparate costs, which might vary with respect to precision, accuracy, and scope. A standardized approach for IAS costing would better enable the comparison of cost estimates between taxa and across studies by controlling for surveying and scaling inconsistencies. This study develops a simple, survey-based approach to generate economic cost estimates for non-native freshwater invasive species (FIS) in Great Britain. The approach scales an average cost for each species by a ratio of management effort, thereby estimating the actual, annual expenditures incurred by a variety of stakeholders. The Great Britain-wide cost of controlling FIS is estimated to be approximately ￡26.5 million year⁻¹; however, the costs of control could total ￡43.5 million year⁻¹ if management efforts were undertaken at all FIS infested locations. Cost estimates are highest for Canadian pondweed (Elodea canadensis), a particularly widespread species, and for the zebra mussel (Dreissena polymorpha), which adversely impacts both industrial water users and boaters. This assessment of the relative economic impacts between species provides policy-makers with a monetary basis for rank-ordering species’ economic impacts and prioritizing management efforts. In addition, the cost assessment approach developed in this study could serve as a model for IAS economic impact assessments elsewhere.     

The Great Lakes in East Africa: biological conservation considerations for species flocks

The three largest water bodies of East Africa, Lake Victoria, Tanganyika, and Malawi contain an estimated number of 2,000 endemic cichlid fish species, in addition, to a mostly uncounted wealth of invertebrates. While the terrestrial diversity is reasonably well protected, as economic and touristic interests coincide with biological conservation strategies, this is not the case for most African lakes and rivers. Nonetheless, it must be promoted that these aquatic ecosystems also deserve protection. Conservation strategies for aquatic biota have so far been the same as for terrestrial environments, i.e., by declaring biodiversity hotspots national parks. Such parks also contain rivers and lake shores. Here, I argue that it seems questionable that this strategy will work, given strong micro-geographic structure of the species flocks and the great degree of local endemism. I suggest a novel strategy for protecting African Lake communities that accounts for local endemism, derived from recent molecular phylogenetic and phylogeographic studies on East African cichlid fishes. While connectivity is the major problem for terrestrial and marine national parks, to ensure a large enough effective population size of the protected animals, this is not the case in most taxa of African rivers and lakes, where local endemism prevails. For example, most littoral cichlid species are subdivided into numerous distinct “color morphs” with restricted distribution, and unlike marine fishes with planktonic larvae display brood care with small offspring numbers. It is argued that the establishment of “micro-scale protected areas,” a large number of small stretches of strictly protected coast line, each only some hundreds of meters long, is likely to work best to preserve the littoral communities in African lakes. Such protected zones can sustain a reasonably effective population size of littoral species, serve as protected spawning ground or nursery area for pelagic species, and at the same time re-seed neighboring populations that are exploited continuously. As long-term stability of littoral fishing grounds is in the immediate interest of village communities, such small protected areas should be managed and controlled by the local communities themselves, and supervised by governmental institutions.     

Global ecological impacts of invasive species in aquatic ecosystems

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta‐analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far‐reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems.     

Non-indigenous species in the Great Lakes: were colonization and damage to ecosystem health predictable?

The Great Lakes ecosystem is home to at least 139 non-indigenous species of fauna and flora which have become established following invasions or intentional introductions. About ten percent of the exotic species have caused economic or ecological damage to the system. A sample of this group is reviewed to determine if ecological concepts are useful in helping to predict colonization and impacts to ecosystem health. Successful colonization by most of the species reviewed was predictable from habitat requirements and behaviour. Ecosystem disturbance was a factor in the success of some of the colonists but was not an overriding ecological requirement. Perturbations to ecosystem health are more difficult to predict and in most cases were not readily apparent from knowledge about the ecology of invaders or native communities. The main damage to ecosystem health by the species reviewed resulted from competition, predation and habitat modification. Difficulties in predicting both invasions and damage from successful colonists point to the need to prevent non-indigenous species from reaching the Great Lakes basin.     

Recent mass invasion of the North American Great Lakes by Ponto-Caspian species

The North American Great Lakes have been invaded and dramatically altered by more than 145 alien species. Many invasions have occurred during the past few decades because of the release of Eurasian ballast water from transoceanic ships. Current regulations require ships to exchange foreign ballast with highly saline water before entering the Great Lakes; this procedure should prevent colonization by strictly freshwater species, but species with broad salinity tolerance might survive transport in exchanged water. A recent series of invasions by euryhaline organisms from the Black and Caspian Seas region signals a new phase in the transformation of the Great Lakes - one that supports the concept of an 'invasional meltdown'.     

Potential for seed bank formation in seven Great Lakes sand dune species

Seeds of seven dune species were collected from sand dunes of Lakes Erie and Huron and buried to various depths in a natural sand dune habitat along Lake Huron. The seed samples were then retrieved after varying lengths of time and examined for their germinability and dormancy. Results showed that buried seeds remained viable for at least 2.5 years and had the potential to form a persistent seed bank. Seed banks were larger and longer lasting at greater depths of burial than those at shallow burial depths. The results suggested that failure to verify the existence of effective seed banks in previous studies may be due to insufficient number of samples, shallow sample depth, local population variations, and fruiting events. Several species also possessed a temporary, aboveground seed reserve formed by retention of a small proportion of viable seeds on the previous year's inflorescences. In some species, seeds retained aboveground were dormant and thus capable of forming a persistent seed bank when they entered the soil.     

Differential invasion success in aquatic invasive species: the role of within- and among-population genetic diversity

Despite a well-developed theoretical basis for the role of genetic diversity in the colonization process, contemporary investigations of genetic diversity in biological invasions have downplayed its importance. Observed reductions in genetic diversity have been argued to have a limited effect on the success of establishment and impact based on empirical studies; however, those studies rarely include assessment of failed or comparatively less-successful biological invasions. We address this gap by comparing genetic diversity at microsatellite loci for taxonomically and geographically paired aquatic invasive species. Our four species pairs contain one highly successful and one less-successful invasive species (Gobies: Neogobius melanostomus, Proterorhinus semilunaris; waterfleas: Bythotrephes longimanus, Cercopagis pengoi; oysters: Crassostrea gigas, Crassostrea virginica; tunicates: Bortylloides violaceous, Botryllus schlosseri). We genotyped 2717 individuals across all species from multiple locations in multiple years and explicitly test whether genetic diversity is lower for less-successful biological invaders within each species pair. We demonstrate that, for gobies and tunicates, reduced allelic diversity is associated with lower success of invasion. We also found that less-successful invasive species tend to have greater divergence among populations. This suggests that intraspecific hybridization may be acting to convert among-population variation to within-population variation for highly successful invasive species and buffering any loss of diversity. While our findings highlight the species-specific nature of the effects of genetic diversity on invasion success, they do support the use of genetic diversity information in the management of current species invasions and in the risk assessment of potential future invaders.     

Genetic diversity in three invasive clonal aquatic species in New Zealand

Background

Elodea canadensis, Egeria densa and Lagarosiphon major are dioecious clonal species which are invasive in New Zealand and other regions. Unlike many other invasive species, the genetic variation in New Zealand is very limited. Clonal reproduction is often considered an evolutionary dead end, even though a certain amount of genetic divergence may arise due to somatic mutations. The successful growth and establishment of invasive clonal species may be explained not by adaptability but by pre-existing ecological traits that prove advantageous in the new environment. We studied the genetic diversity and population structure in the North Island of New Zealand using AFLPs and related the findings to the number of introductions and the evolution that has occurred in the introduced area.

Results

Low levels of genetic diversity were found in all three species and appeared to be due to highly homogeneous founding gene pools. Elodea canadensis was introduced in 1868, and its populations showed more genetic structure than those of the more recently introduced of E. densa (1946) and L. major (1950). Elodea canadensis and L. major, however, had similar phylogeographic patterns, in spite of the difference in time since introduction.

Conclusions

The presence of a certain level of geographically correlated genetic structure in the absence of sexual reproduction, and in spite of random human dispersal of vegetative propagules, can be reasonably attributed to post-dispersal somatic mutations. Direct evidence of such evolutionary events is, however, still insufficient.     

From family to nation: Organizing the Mississippi movement

John Dittmer, LOCAL PEOPLE: THE STRUGGLE FOR CIVIL RIGHTS IN MISSISSIPPI, Urbana, IL: University of Illinois Press, 1995, 530 pp., $29.95.

Charles M. Payne, I'VE GOT THE LIGHT OF FREEDOM: THE ORGANIZING TRADITION AND THE MISSISSIPPI FREEDOM STRUGGLE, Berkeley, CA: University of California Press, 1995, xiv + 525 pp, $28.00.