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.     

Linking fish population dynamics to habitat conditions: insights from the application of a process-oriented approach to several Great Lakes species

One of the major challenges facing fishery scientists and managers today is determining how fish populations are influenced by habitat conditions. Many approaches have been explored to address this challenge, all of which involve modeling at one level or another. In this paper, we explore a process-oriented model approach whereby the critical population processes of birth and death rates are explicitly linked to habitat conditions. Application of this approach to five species of Great Lakes fishes including: walleye (Sander vitreus), lake trout (Salvelinus namaycush), smallmouth bass (Micropterus dolomieu), yellow perch (Perca flavescens), and rainbow trout (Onchorynchus mykiss), yielded a number of insights into the modeling process. One of the foremost insights is that processes determining movement and transport of fish are critical components of such models since these processes largely determine the habitats fish occupy. Because of the importance of fish location, an individual-based model appears to be a nearly inescapable modeling requirement. There is, however, a paucity of field-based data directly relating birth, death, and movement rates to habitat conditions experienced by individual fish. There is also a paucity of habitat information at a fine temporal and spatial scale for many important habitat variables. Finally, the general occurrence of strong ontogenetic changes in the response of different life stages to habitat conditions emphasizes the need for a modeling approach that considers all life stages in an integrated fashion.     

Description of a new species of myxozoan from Notropis hudsonius in the Great Lakes region of Canada

Myxobolus burti n. sp. is described from striated muscle of Notropis hudsonius (Cyprinidae) collected from localities in Lake Superior, Lake Michigan, Lake Huron, Lake St. Claire, Lake Erie, Lake Ontario, and the St. Lawrence River. Myxobolus burti is intracellular, forming thin-walled, cigar-shaped plasmodia within striated muscle cells of the body. It exhibits disporoblastic, oval spores, approximately 10 μm long, 8 μm wide, and 6 μm thick, with a length-to-width ratio of 1.3 ∶ 1. They resemble most closely those of Myxobolus bellus Kudo, 1934, and Myxobolus mutabilis Kudo, 1934, but have polar capsules unequal in length (5.3 vs. 4.7 μm) and oblique filament coils. The new parasite was not encountered in routine examination of other small-bodied fishes at collection locations and thus looks to be specific to N. hudsonius. A comparative plate of similar species encountered during surveys of these fishes in the northeast is presented to contrast the uniqueness of the new species.     

Space–time patterns during the establishment of a nonindigenous species

Increasing rates of global trade and travel have the invariable consequence of an increase in the likelihood of nonindigenous species arrival, and some new arrivals are successful in establishing themselves. Quantifying the pattern of establishment of nonindigenous species across both spatial and temporal scales is paramount in early detection efforts, yet very difficult to accomplish. Previous work in epidemiology has proposed methods for assessing the space–time properties of emerging infectious diseases by quantifying the degree of space–time clustering between individual cases. I tested the applicability of one such method commonly used in epidemiology, the Knox test for space–time interaction, to analyze rare abundance data from an isolated, newly-establishing gypsy moth, Lymantria dispar, population in Minnesota, USA, and incorporated a bootstrap approach to quantify the space–time pattern in a random process that can be used to compare with results from empirical data. The use of the Knox test in assessing the establishment phase of biological invasions could potentially serve as an early warning system against new invaders, particularly for those with a known history of a repeated number of arrivals.     

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'.     

Dispersal strategies, secondary range expansion and invasion genetics of the nonindigenous round goby, Neogobius melanostomus, in Great Lakes tributaries

Dispersal strategies are important mechanisms underlying the spatial distribution and colonizing ability of all mobile species. In the current study, we use highly polymorphic microsatellite markers to evaluate local dispersal and colonization dynamics of the round goby (Neogobius melanostomus), an aquatic invader expanding its range from lake to river environments in its introduced North American range. Genetic structure, genotype assignment and genetic diversity were compared among 1262 round gobies from 20 river and four lake sites in three Great Lakes tributaries. Our results indicate that a combination of short-distance diffusion and long-distance dispersal, collectively referred to as 'stratified dispersal', is facilitating river colonization. Colonization proceeded upstream yearly (approximately 500 m/year; 2005-2009) in one of two temporal replicates while genetic structure was temporally stable. Contiguous dispersal from the lake was observed in all three rivers with a substantial portion of river fish (7.3%) identified as migrants. Genotype assignment indicated a separate introduction occurred upstream of the invasion front in one river. Genetic diversity was similar and relatively high among lake and recently colonized river populations, indicating that founder effects are mitigated through a dual-dispersal strategy. The remarkable success of round goby as an aquatic invader stresses the need for better diffusion models of secondary range expansion for presumably sessile invasive species.     

A multi-modeling approach to evaluating climate and land use change impacts in a Great Lakes River Basin

River ecosystems are driven by linked physical, chemical, and biological subsystems, which operate over different temporal and spatial domains. This complexity increases uncertainty in ecological forecasts, and impedes preparation for the ecological consequences of climate change. We describe a recently developed “multi-modeling” system for ecological forecasting in a 7600 km² watershed in the North American Great Lakes Basin. Using a series of linked land cover, climate, hydrologic, hydraulic, thermal, loading, and biological response models, we examined how changes in both land cover and climate may interact to shape the habitat suitability of river segments for common sport fishes and alter patterns of biological integrity. In scenario-based modeling, both climate and land use change altered multiple ecosystem properties. Because water temperature has a controlling influence on species distributions, sport fishes were overall more sensitive to climate change than to land cover change. However, community-based biological integrity metrics were more sensitive to land use change than climate change; as were nutrient export rates. We discuss the implications of this result for regional preparations for climate change adaptation, and the extent to which the result may be constrained by our modeling methodology.     

Exotic plant species of the St Lawrence River wetlands: a spatial and historical analysis

Aim To evaluate the importance (number of species, plant cover) of the exotic flora in seven well‐defined sectors of one of the most important transportation waterways in North America. To determine the impact of exotic species on wetland plant diversity and reconstruct the spread of some invasive species. Location St Lawrence River, southern Québec. Methods The exotic flora (vascular plants) of wetlands bordering the St Lawrence River was studied using 713 sampling stations (25 m²) along a 560‐km long corridor. Results Exotic species represent 13.7% of the vascular flora of the St Lawrence wetlands. The relative plant cover occupied by exotic species is high in some of the fluvial sectors (42–44%), but low (6–10%) in the estuarine sectors. Wetlands (marshes) surrounding islands were particularly susceptible to invasion by exotic plants. Historical, abiotic and landscape factors may explain the differences observed between sites. Purple loosestrife (Lythrum salicaria L.) is the most common exotic species of the St Lawrence wetlands, but other species, namely flowering‐rush (Butomus umbellatus L.) and reed canary grass (Phalaris arundinacea L.) are much more invasive. There is no linear relationship between the exotic species cover and the diversity of wetland plants; low diversity sites can be dominated by either exotic or native plant species. In the other sites, exotic species generally have little impact on plant communities and can contribute to increase diversity. Common reed (Phragmites australis (Cav.) Trin. ex Steudel) and reed canary grass, both considered as exotic species in this study, clearly have a stronger impact on plant diversity than flowering‐rush and purple loosestrife. Main conclusions This study shows that the global impact of an invader cannot be adequately evaluated with only a few highly invaded sites. While nationwide strategies have been developed to control exotic species, large surveys are essential to adapt them to regional particularities.     

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.     

Potential effects on zooplankton from species shifts in planktivorous mussels: a field experiment in the St Lawrence River

1. Suspension feeding by bivalves exceeds that by other planktivores in many North American rivers, and food webs may be altered substantially by differences in feeding patterns between native unionid mussels and invading dreissenid mussels. 2. We conducted an experiment comparing zooplanktivory by one unionid and two dreissenid species that addressed several primary questions. Is benthic planktivory important in this river? Has this linkage been altered substantially by dreissenids? Do the two dreissenid species differ in planktivory, and is this ecologically important if quagga mussels extend their geographical range? 3. Our 12‐day experiment consisted of controls (no mussels) and treatments with unionid (Elliptio complanata), quagga (Dreissena bugensis) or zebra (D. polymorpha) mussels in 3500‐L, 80‐μm mesh enclosures placed in a slackwater area of the St Lawrence River. 4. The density of the most abundant calanoid copepod Eurytemora affinis increased in the presence of dreissenids, probably as an indirect food web response. By day 12, a cumulative effect was shown by the most overwhelmingly abundant rotifer, Polyarthra, whose density declined dramatically in dreissenid enclosures compared with control and unionid enclosures. Rotifer densities in unionid enclosures were not different from controls, nor were dreissenid treatments different from each other. The effects on rotifers were probably from predation, as Chl‐a did not vary among treatments. 5. We conclude that benthic‐pelagic coupling via planktivory is important in slackwater areas. Dreissenids have strengthened this linkage, but range extension of quaggas should not appreciably alter effects produced by a similar biomass of zebra mussels.     

Waiting for invasions: a framework for the arrival of nonindigenous species

The process of nonindigenous species (NIS) arrival has received limited theoretical consideration despite importance in predicting and preventing the establishment of NIS. We formulate a mechanistically based hierarchical model of NIS arrival and demonstrate simplifications leading to a marginal distribution of the number of surviving introduced individuals from parameters of survival probability and propagule pressure. The marginal distribution is extended as a stochastic process from which establishment emerges with a waiting time distribution. This provides a probability of NIS establishment within a specified period and may be useful for identifying patterns of successful invaders. However, estimates of both the propagule pressure and the individual survival probability are rarely available for NIS, making estimates of the probability of establishment difficult. Alternatively, researchers are able to measure proportional estimates of propagule pressure through models of NIS transport, such as gravity models, or of survival probability through habitat-matching indexes measuring the similarity between potentially occupied and native NIS ranges. Therefore, we formulate the relative waiting time between two locations and the probability of one location being invaded before the other.     

Secondary invasion of the round goby into high diversity Great Lakes tributaries and species at risk hotspots: potential new concerns for endangered freshwater species

The round goby (Neogobius melanostomus) first invaded North America in 1990 when it was discovered in the St. Clair River. Despite more than 15 years of potential invasion, many Great Lakes’ lotic systems remained uninvaded. Recently, we captured the round goby from several Great Lakes tributaries known as species-at-risk hotspots. With a combination of field sampling of round gobies and literature review of the impact of round gobies on native taxa, we assess the potential impacts of the secondary invasion to native species using three mechanisms: competition; predation; and indirect impacts from the loss of obligate mussel hosts. We estimate that 89% (17/19) of benthic fishes and 17% (6/36) of mussels that occur in these systems are either known or suspected to be impacted by the secondary invasion of round goby. In particular, we note that the distribution of potential impacts of round goby invasion was largely associated with species with a conservation designation, including seven endangered species (1 fish, 6 mussels). As these recent captures of round goby represent novel occurrences in high diversity watersheds, understanding the potential impacts of secondary invasion to native biota is fundamental to prevent species declines and to allow early mitigation.     

The response of zooplankton and phytoplankton from the North American Great Lakes to filtration

Filtration of ballast water was investigated as a means of minimizing the introduction of nonindigenous zooplankton and phytoplankton by ships visiting the North American Great Lakes-St. Lawrence Seaway system (GLSLSS). An automatic backwash screen filtration (ABSF) system with nominal filtration options of 25, 50 or 100 μm was mounted on the deck of an operating Seaway-sized dry bulk carrier, the MV Algonorth. Water was pumped through the ABSF with a deck mounted pump at 341 m³ hr⁻¹ during routine ship operations in the GLSLSS, and effectiveness of the various screen pore sizes at removing taxonomic categories of zooplankton and phytoplankton was measured using matched treatment and control ballast tanks. The smallest pore sizes (25 and 50 μm) performed better than the 100 μm pore size at removing biological material. There was no difference in the filtration efficiency of the 25 and 50 μm screens relative to macro- or microzooplankton in these tests, but this result was probably due to low densities of macrozooplankton, and soft-bodied (aloricate) characteristics of the microzooplankton present. The 25 and 50 μm pore sizes were subjected to more controlled tests on board a stationary barge platform equipped with triplicate 700 L catchment bins moored in Duluth Harbor of Lake Superior. In these tests, filter pore size, organism size and rigidity influenced zooplankton removal efficiency by the ABSF. The 25 μm screen reduced both macrozooplankton and microzooplankton significantly more than the 50 μm screen. Zooplankton width was more determinative of filtration performance than length, and both filters removed loricate species of rotifers significantly more efficiently than aloricate species of the same length and width size classes. The 25 and 50 μm ABSF also significantly reduced algal densities, with the exception of colonial and filamentous green algae (50 μm only). Filter efficiency relative to algal particles was influenced by filter pore size, organism morphology and structure, and intake density, while algal particle size was not determinative. This research provides compelling evidence that 25 or 50 μm filtration is a potentially powerful means of reducing densities of organisms discharged by ships operating in the Great Lakes but an additional treatment step would be necessary to effectively minimize risk and meet the International Maritime Organization's discharge standards associated with organisms of all sizes in the water column.     

Restoration of rapids habitat in a Great Lakes connecting channel,the St. Marys River,Michigan

Aquatic habitat has been extensively altered throughout the Laurentian Great Lakes to increase navigation connectivity. In particular, the St. Marys River, a Great Lakes connecting channel, lost >50% of its historic rapids habitat over the past century. In 2016, the natural flow was restored to the Little Rapids area of the St. Marys River. The goal of our study was to evaluate physical and ecological responses to the restoration of the Little Rapids area. Extensive habitat and biological data were collected prior to restoration (2013 and 2014), and after restoration (2017 and 2018). Measured parameters included total suspended solids, current velocity, benthic macroinvertebrates, and larval, juvenile, and adult fishes. Total suspended solids stayed low (<4 mg/L) following restoration, with the exception of a single construction‐related event. Pre‐restoration data indicated that all measured velocities were below the target flow rate of 0.24 m/s, whereas 70% of the measured habitat was above the target flow post‐restoration. Abundance and richness of benthic macroinvertebrates were reduced following restoration (>90% reduction). We observed a 45% increase in richness of larval fish 2 years after restoration and a 131% increase in catch per unit effort. For adult fishes, the proportion of individuals with a preference for fast‐moving waters increased from 1.5 to 45% in the restored area, and from 7 to 15% upstream of the restored area; a similar response was observed for lithophilic spawners. The physical and biological conditions of the Little Rapids improved and resembled conditions typical of rapids habitat extent in other areas of the river and other systems.     

The robustness of geometric morphometrics in testing the morphological equivalence hypothesis among cichlid species from East African Great Lakes

As part of an ongoing broader investigation into whether similar morphologies among cichlid species from East African Great Lakes are due to common ancestry or convergent evolution, we used geometric morphometrics to quantitatively test the hyphothesis that Lake Tanganyika's Tropheus spp. is morphologically similar to Lake Malawi's Pseudotropheus spp. and Labeotropheus fuelleborni . Landmark-based techniques were used to investigate body shape variation and the results revealed highly significant differences among these genera with the exception of L. fuelleborni vs. Pseudotropheus ( Tropheops ) spp. An ordination of all specimens along the first two canonical variate (CV) axes indicated clear interlake separation, especially along the CV I axis, whereby Tropheus moori showed a dorsoventral expansion of the mid body section with a steeper forehead relative to Lake Malawi species. Pseudotropheus ( Maylandia ) zebra was discriminated from the rest primarily on the basis of its terminally positioned gape, which differed from the inferior subterminal gape that was prevalent in the other species. Thus, the present study reveals a pattern of parallel evolution between L. fuelleborni and Pseudotropheus ( Tropheops ) spp., but does not support an interlake convergent evolution hypothesis. Contrary to previous researchers' assertions, these results emphasize the robustness of geometric morphometrics when testing morphological equivalence hypothesis among cichlid species.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 1–9.     

An heuristic model of transformations of the aquatic ecosystems of the Great Lakes-St. Lawrence River Basin

Seminal works of the 1960s on open systems by Arthur Koestler, Ramon Margalef and Richard Vollenweider are interrelated here, with many other works on the science of aquatic ecosystems, in a heuristic model of selected manifestations and transformations of the aquatic ecosystems of the rivers and lakes of the Great Lakes — St. Lawrence River Basin, here perceived as a single loosely-integrated ecosystem. Our empirical model is in the form of a tentative generalization of a set of features and events — large-scale with respect to time and space — in these aquatic ecosystems, with some suggestions of recent relevant theoretical concepts from post-normal studies of complex phenomena. Our work may be a step toward a more comprehensive, bioregional, heuristic model of the Basin to include the aquatic and terrestrial, as well as the natural and cultural ecosystemic features.