Metazoan Parasites of Introduced Round and Tubenose Gobies in the Great Lakes: Support for the “Enemy Release Hypothesis”

Recent invasion theory has hypothesized that newly established exotic species may initially be free of their native parasites, augmenting their population success. Others have hypothesized that invaders may introduce exotic parasites to native species and/or may become hosts to native parasites in their new habitats. Our study analyzed the parasites of two exotic Eurasian gobies that were detected in the Great Lakes in 1990: the round goby Apollonia melanostoma and the tubenose goby Proterorhinus semilunaris. We compared our results from the central region of their introduced ranges in Lakes Huron, St. Clair, and Erie with other studies in the Great Lakes over the past decade, as well as Eurasian native and nonindigenous habitats. Results showed that goby-specific metazoan parasites were absent in the Great Lakes, and all but one species were represented only as larvae, suggesting that adult parasites presently are poorly-adapted to the new gobies as hosts. Seven parasitic species are known to infest the tubenose goby in the Great Lakes, including our new finding of the acanthocephalan Southwellina hispida, and all are rare. We provide the first findings of four parasite species in the round goby and clarified two others, totaling 22 in the Great Lakes—with most being rare. In contrast, 72 round goby parasites occur in the Black Sea region. Trematodes are the most common parasitic group of the round goby in the Great Lakes, as in their native Black Sea range and Baltic Sea introduction. Holarctic trematode Diplostomum spathaceum larvae, which are one of two widely distributed species shared with Eurasia, were found in round goby eyes from all Great Lakes localities except Lake Huron proper. Our study and others reveal no overall increases in parasitism of the invasive gobies over the past decade after their establishment in the Great Lakes. In conclusion, the parasite “load” on the invasive gobies appears relatively low in comparison with their native habitats, lending support to the “enemy release hypothesis.”     

Parasites of Lake Trout,Salvelinus namaycush,from the Great Lakes: a Review of the Literature 1874–1994

A comparative summary of the literature on parasites reported from lake trout, Salvelinus namaycush, from the Great Lakes is presented. Twenty-four studies on parasites infecting lake trout in these lakes have been published since 1874 with 32 parasite species (1 Monogenea, 3 Digenea, 9 Cestoidea, 6 Nematoda, 6 Acanthocephala, 1 Annelida, 5 Crustacea, 1 Fungi) being reported. The most common parasites are Eubothrium salvelini and Echinorhynchus salmonis in the intestine, and Cystidicola stigmatura in the swim bladder. Most of the parasites occur in the gastrointestinal tract. Twenty (80%) of the 25 helminth species reported from lake trout are represented by adults. The dominance of these helminth species is attributable to the position of lake trout in the food web of the Great Lakes as top predators. Parasitological data from splake, Salvelinus namaycush X Salvelinus fontinalis, from Lake Huron are also presented.     

Revision of the needle-shaped Fragilaria species (Fragilariaceae,Bacillariophyta) in the Laurentian Great Lakes (United States of America,Canada)

We investigated several common, needle-shaped diatoms to better characterize the taxonomy of the genus Fragilaria in the Laurentian Great Lakes. We conducted diatom morphometric analysis facilitated by SEM and LM imaging on samples collected as a part of the USEPA’s long-term biological monitoring program. We resolved several decades-long taxonomic problems in the Great Lakes. The results indicated that previous records of species formerly belonging to the genus Synedra, such as S. (Fragilaria) ostenfeldii, S. (Fragilaria) radians, and S. filiformis, should be corrected as these species likely do not occur in the Great Lakes. Valve morphology confirmed the presence of four previously undescribed species: Fragilaria andreseniana sp. nov., Fragilaria stoermeriana sp. nov., Fragilaria limnetica sp. nov., and Fragilaria michiganensis sp. nov. The morphology of several other Fragilaria taxa in the Great Lakes was examined, including a teratologic taxon (Fragilaria sp. 1), Fragilaria crotonensis, Fragilaria grunowii, and a taxon showing morphological affinity to Fragilaria lemanensis (reported here as Fragilaria cf. lemanensis). The spatial and seasonal distributions of these needle-shaped taxa were also analyzed and discussed.     

Status of the major aquaculture carps of China in the Laurentian Great Lakes Basin

There is concern of economic and environmental damage occuring if any of the four major aquacultured carp species of China, black carp Mylopharyngodon piceus, bighead carp Hypophthalmichthys nobilis, silver carp H. molitrix, or grass carp Ctenopharyngodon idella, were to establish in the Laurentian Great Lakes. All four are reproducing in the Mississippi River Basin. We review the status of these fishes in relation to the Great Lakes and their proximity to pathways into the Great Lakes, based on captures and collections of eggs and larvae. No black carp have been captured in the Great Lakes Basin. One silver carp and one bighead carp were captured within the Chicago Area Waterway System, on the Great Lakes side of electric barriers designed to keep carp from entering the Great Lakes from the greater Mississippi River Basin. Three bighead carp were captured in Lake Erie, none later than the year 2000. By December 2019, at least 650 grass carps had been captured in the Great Lakes Basin, most in western Lake Erie, but none in Lake Superior. Grass carp reproduction has been documented in the Sandusky and Maumee rivers in Ohio, tributaries of Lake Erie. We also discuss environmental DNA (eDNA) results as an early detection and monitoring tool for bighead and silver carps. Detection of eDNA does not necessarily indicate presence of live fish, but bigheaded carp eDNA has been detected on the Great Lakes side of the barriers and in a small proportion of samples from the western basin of Lake Erie.     

Influence of river plumes on the distribution and composition of nearshore Lake Michigan fishes

River plumes form in coastal areas where tributaries mix with their receiving waters. Plume waters are enriched with terrestrial-derived nutrients from their watersheds creating hotspots of biological productivity. The biological importance of plumes scales with the size and persistence of the plume; therefore, large, persistent plumes are more important than small, transient plumes. To date, most studies of plumes have focused on assimilation of terrestrial-derived energy by aquatic species or lower-level food web effects, primarily in marine systems. Few studies have described fish communities near plume habitats and compared them to non-plume areas, especially for the numerous small plumes in the Laurentian Great Lakes. Here we demonstrate that small plumes in the main basin of Lake Michigan enhance local primary productivity and influence distribution and abundance of nearshore Great Lakes fishes. We found that plume fish communities were relatively depauperate and did not support higher biological diversity of fishes compared to non-plume areas. However, individual species including rainbow smelt Osmerus mordax, spottail shiner Notropis hudsonius, and white sucker Catostomus commersonii were more abundant around plumes. Our results demonstrate that small plumes in the main basin of Lake Michigan support highly localized hotspots of biological productivity and fish abundance, primarily within 2?km of river mouths.     

Brachionus leydigii (Monogononta: Ploima) reported from the western basin of Lake Erie

Several species of non-indigenous planktonic invertebrates have historically been introduced to the Laurentian Great Lakes. Previous introductions of non-indigenous planktonic invertebrates to the Great Lakes have been crustacean zooplankton, specifically Cladocera and Copepoda. This report documents the first known occurrence of Brachionus leydigii var. tridentatus (Zernov, 1901) in Lake Erie and possibly the first detection of a non-indigenous rotifer species in the Laurentian Great Lakes. The specimen was collected from a U.S. EPA monitoring station in the western basin of Lake Erie on April 4, 2016.     

An exploration of Lake Ontario’s offshore epibenthic cladoceran community

Pelagic cladocerans are relatively well studied within the Great Lakes. However, little is known about cladocerans that more closely interact with benthic substrate. In 2018, 26 sites in Lake Ontario were sampled with benthic grabs to collect meiofauna. The stations ranged in depth from 0.1 to 184 m with sand, silt or hard bottom substrate. Epibenthic cladocerans were recovered from all 12 sites shallower than 20 m, five of eight sites between 20 and 40 m and one (63 m deep) of six sites deeper than 40 m. The density of epibenthic cladocerans were at most 12% of the density of harpacticoid copepods (shallow hard bottom sites). The Lake Ontario epibenthic cladoceran community was represented by 16 species from 8 different genera although one of these species, Illyocryptus cf. sordidus, is likely represented by two cryptic species. The most widespread species was identified as Alona sibirica (Sinev, 2020), a newly described species from Russia formerly of the Alona cf. affinis (Leydig, 1860) species complex. The Great Lakes specimens were 98.3% similar genetically to sequences assigned to A. cf. sibirica. Among substrates, sand had the highest densities (1662 m² in 0–20 m) and species richness (10) of epibenthic Cladocera. After cladocerans were identified, the cytochrome oxidase I (COI) gene of 37 specimens were successfully sequenced through the Barcode of Life (BOLD). Six specimens were assigned to three previously existing Barcode Identification Numbers (BINs), the remaining 31 specimens were assigned to eight new BINs.     

Diatoms from the Great Lakes. I. Rare or Poorly known Species of the Genera Diploneis,Oestrupia and Stauroneis

Eleven taxa of diatoms occurring in the Great Lakes which have not been adequately treated in the North American literature are briefly described and figured. Taxa treated include 4 species of Diploneis, 1 species and one variety of Oestrupia, and 1 species and 4 infraspecific taxa of Stauroneis. The presently known distribution of these entities in the Great Lakes and their ecological and distributional affinities are discussed.     

The establishment of the nuisance cyanobacteria Lyngbya wollei in Lake St. Clair and its potential to harbor fecal indicator bacteria

Lyngbya wollei is a filamentous cyanobacterium which forms large nuisance mats and has infested eastern and southeastern U.S. Lakes and reservoirs for over 100 years. Lyngbya was recently identified in the Great Lakes system in the St. Lawrence River, and Western Lake Erie. Here we report on large deposits of L. wollei washing onshore at a popular recreational beach in Lake Saint Clair, part of the Great Lakes system. The amount of L. wollei deposited on shore was quantified and evaluated for the presence of fecal indicator bacteria (FIB). High concentrations of Escherichia coli, enterococci and Clostridium perfringens were found in the L. wollei in nearshore waters. The densities of E. coli (MPN), enterococci (MPN) and C. perfringens (CFU) attached to L. wollei averaged 3.5, 3.2 and 3.2 log/g, respectively. In contrast, nearshore waters contained nearly 10 times less FIB, averaging 2.6, 2.4 and 2.6 log/100 ml of E. coli (MPN), enterococci (MPN) and C. perfringens (CFU), respectively. DNA fingerprint analysis was used to examine the population structure of E. coli isolates obtained from L. wollei mats. The L. wollei-borne E. coli strains were genetically diverse, suggesting a causal relationship between E. coli and L. wollei. Results from this study indicate that in addition to the macroalga such as Cladophora, cyanobacteria like L. wollei also harbor FIB, potentially impacting water quality and human health in the Great Lakes.     

Predicting the potential distribution of the non-native Red Swamp Crayfish Procambarus clarkii in the Laurentian Great Lakes

The ongoing threat of introduction of invasive species, including crayfish, to the Laurentian Great Lakes has motivated the development of predictive models to inform where these invaders are likely to establish. Our study is among the first to apply regional freshwater-specific GIS layers to species occurrence data to predict ecosystem suitability to invasions, specifically for the red swamp crayfish, Procambarus clarkii, in the Great Lakes. We combined a database of crayfish species occurrences with the Great Lakes Aquatic Habitat Framework (GLAHF) GIS layers to model habitats suitable to invasion by P. clarkii using boosted regression trees and physiological information for this species. We developed a model of all suitable crayfish habitat across the Great Lakes, then constrained this habitat to areas anticipated to be suitable for P. clarkii based on known physiological limitations of this species. Specifically, P. clarkii requires a minimum temperature of 15?°C for copulation and oviposition, with peak reproduction occurring at temperatures of 20–23?°C. We identified 2% of the Great Lakes as suitable for P. clarkii establishment and 0.88% as optimal for this crayfish, primarily located on the southern coastlines of lakes Michigan and Erie and shallow bays including Saginaw Bay (Lake Huron), Green Bay (Lake Michigan), and Henderson Bay (Lake Ontario). These predictions of where P. clarkii is likely to establish populations can be used to identify areas where education, outreach, compliance, and law enforcement efforts should seek to prevent new introductions of this crayfish and help prioritize locations for surveillance to detect newly established populations.     

First report of tubenose goby (Proterorhinus semilunaris) in the eastern basin of Lake Erie

The tubenose goby (Proterorhinus semilunaris), native to the Ponto-Caspian region, was first discovered in the Laurentian Great Lakes in 1990 after it was introduced through ballast water discharge. Compared with Neogobius melanostomus, another exotic gobiid from the Ponto-Caspian, colonization of the Great Lakes by P. semilunaris has been slow, with reports of the species being largely confined to the Huron-Erie Corridor (HEC) and western portions of Lake Erie and Lake Superior. This is the first report of P. semilunaris in the Great Lakes east of the western basin of Lake Erie. Between 28 June and 27 July, 2012, 176 P. semilunaris were collected from shallow (< 1.2 m) water of Marina Lake, a 40 ha embayment in Presque Isle State Park (Erie, PA). The large number of P. semilunaris collected at the site and the presence of individuals as small as 17 mm total length suggest an established population. However, the mechanism by which P. semilunaris was introduced to Presque Isle Bay is not clear.     

Daphnia lumholtzi Sars a new alien species in Lake St. Clair

Daphnia lumholtzi Sars, an exotic tropical/subtropical cladoceran from Australia, southeast Asia, and Africa, was newly found in Lake St. Clair in a vertical tow sample taken at 3 m depth on 25 July 2007. The species was previously found in 1990/1991 in some reservoirs in the southern United States from where it colonized many waters north to the Great Lakes. In 1999, it was found in Lake Erie. This cladoceran had a density of 117 individuals/m³ when we collected it from Lake St. Clair and it was represented by both females (95.12 %) and males (4.88 %). It seems that D. lumholtzi will continue to expand its distribution area in the Great Lakes.     

Early observations on an emerging Great Lakes invader Hemimysis anomala in Lake Ontario

Hemimysis anomala, a Ponto-Caspian littoral mysid, is an emerging Great Lakes invader that was discovered in Lakes Michigan and Ontario in 2006. Similar to the native mysid Mysis diluviana, Hemimysis exhibits a diel vertical migration pattern but generally inhabits shallower and warmer waters than M. diluviana. Because basic information on the distribution, habitat use, and biology of Hemimysis in the Great Lakes is scarce, the potential for food web disruption by Hemimysis cannot easily be predicted. Preliminary observations indicate widespread invasion of Hemimysis in Lake Ontario. In this study, we confirm the presence of Hemimysis at sites spanning the northern and southern shores of Lake Ontario and the presence of the individuals during winter months. In one horizontal tow in November 2007, over 26,000 individuals were collected with a length range of 4.4 to 9.0 mm and an average caloric density of 611 cal/g wet weight. The most effective methods for sampling Hemimysis were horizontal tows with either a zooplankton net in the water column or a benthic sled near the lake bottom. Although more quantitative data on the life history and distribution of this species is necessary, our preliminary observations support the prediction that the potential for Hemimysis to impact the nearshore food web in Lake Ontario appears high.     

Historical changes and current status of crayfish diversity and distribution in the Laurentian Great Lakes

Despite increasing recognition of the importance of invertebrates, and specifically crayfish, to nearshore food webs in the Laurentian Great Lakes, past and present ecological studies in the Great Lakes have predominantly focused on fishes. Using data from many sources, we provide a summary of crayfish diversity and distribution throughout the Great Lakes from 1882 to 2008 for 1456 locations where crayfish have been surveyed. Sampling effort was greatest in Lake Michigan, followed by lakes Huron, Erie, Superior, and Ontario. A total of 13 crayfish species occur in the lakes, with Lake Erie having the greatest diversity (n = 11) and Lake Superior having the least (n = 5). Five crayfish species are non-native to one or more lakes. Because Orconectes rusticus was the most widely distributed non-native species and is associated with known negative impacts, we assessed its spread throughout the Great Lakes. Although O. rusticus has been found for over 100 years in Lake Erie, its spread there has been relatively slow compared to that in lakes Michigan and Huron, where it has spread most rapidly since the 1990s and 2000, respectively. O. rusticus has been found in both lakes Superior and Ontario for 22 and 37 years, respectively, and has expanded little in either lake. Our broad spatial and temporal assessment of crayfish diversity and distribution provides a baseline for future nearshore ecological studies, and for future management efforts to restore native crayfish and limit non-native introductions and their impact on food web interactions.     

The Invasive New Zealand Mud Snail (Potamopyrgus antipodarum) in Lake Erie

The New Zealand mud snail (Potamopyrgus antipodarum) is an invasive species in Europe, Japan, Australia, and North America. In the western United States it is a species of special concern where population densities in some rivers and streams are very large (∼300,000 per m²) and considerable ecological effects of its presence have been reported. Much less about the effects of this species is known in the Great Lakes, where the snail was found in Lake Ontario and the St. Lawrence River in 1991. Here we report the occurrence of the snail in Lake Erie. Two P. antipodarum were collected in 18 m deep water (sampling range 5–18 m) in Lake Erie off shore of Presque Isle State Park near Erie, Pennsylvania in the summer of 2005 and others were collected off of Sturgeon Point in Lake Erie (sampling range 5–20 m) south of Buffalo, NY and in the central basin of Lake Erie (18 m) in 2006. This finding demonstrates that this species continues to expand its range in the Great Lakes. The range expansion increases the likelihood that it may become established in rivers and streams emptying into the Great Lakes where higher densities and greater ecological damage may result.     

Relative abundance,age structure,and body size in mudpuppy populations in southwestern Ontario

Little is known of mudpuppy (Necturus maculosus) population structure and ecology; some populations in the Great Lakes are thought to be in decline. Mudpuppies are the obligate hosts for the mudpuppy mussel (Simpsonaias ambigua), a species that is endangered in Canada and in many Great Lakes states. We surveyed mudpuppies from the Sydenham River, the only known Canadian locality of the mudpuppy mussel, in order to generate information on relative density, deformity rates and population age/size structure and used this information to compare them to known mudpuppy populations from Great Lakes sites in the Detroit River, Lake St. Clair and Long Point in Lake Erie. Deformity rates were elevated at some sites in the Sydenham River. The relative density of mudpuppies in the Sydenham River was lower than that of other Great Lakes sites and their age was skewed towards younger individuals. Although at lower densities than at other Great Lakes sites, the mudpuppy population in the Sydenham River appears stable and is showing signs of recruitment which bodes well for the future of the mudpuppy mussel population of the river.     

NOTE: First record of natural reproduction by Atlantic salmon (Salmo salar) in the St. Marys River,Michigan

Atlantic salmon (Salmo salar) are native to Lake Ontario; but their populations severely declined by the late 1800s due to human influences. During the early to mid-1900s, Atlantic salmon were stocked throughout the Great Lakes in effort to reestablish them into Lake Ontario and introduce the species into the upper Great Lakes. However, these efforts experienced minimal success. In 1987, Lake Superior State University and the Michigan Department of Natural Resources began stocking Atlantic salmon in the St. Marys River, Michigan, which has resulted in a successful, self-supporting hatchery operation and stable recreational Atlantic salmon fishery. Possibly due to a combination of competition with other salmonid species for spawning habitat, prey selection causing detrimental effects on early life stages and high rates of early mortality syndrome, Atlantic salmon appeared to be severely limited in their ability to naturally reproduce within the upper Great Lakes. In 2012, the first unequivocal documentation of naturally reproduced Atlantic salmon in the St. Marys River was recorded, downstream from the compensation works and parallel to the Soo Locks in Sault Ste. Marie, Michigan.     

Origins of Rainbow Smelt in Lake Ontario

The first rainbow smelt (Osmerus mordax) to enter Lake Ontario were probably migrants from an anadromous strain introduced into New York's Finger Lakes. Since the upper Great Lakes were originally stocked with a landlocked strain from Green Lake, Maine, subsequent migration to Lake Ontario from Lake Erie makes Lake Ontario unique among the Great Lakes in probably having received introductions from two distinct populations.     

Invasive Plant Species in Diked vs. Undiked Great Lakes Wetlands

We compared the standing vegetation, seed banks, and substrate conditions in seven pairs of diked and undiked wetlands near the shores of Lake Michigan and Lake Huron, North America. Our analysis tested the null hypothesis that construction of artificial dikes has no effect on the vulnerability of Great Lakes coastal wetlands to non-native and native invasive species. Both the standing vegetation and seed banks in diked wetlands contained significantly more species and individuals of invasive plants. In addition, diked wetlands exhibited significantly higher levels of organic matter and nutrient levels, and significantly higher average pH. Two pervasive non-native invasive species in the Great Lakes region, Lythrum salicaria (purple loosestrife) and Phalaris arundinacea (reed canary grass) were significantly more abundant in diked wetlands. Typha spp. (cattail) also formed a much higher percent vegetation cover in the diked wetlands. Our results support the view that diking of shoreline wetlands modifies natural hydrologic regimes, leading to nutrient-rich aquatic environments that are vulnerable to invasion. The shallower, more variable water levels in non-diked wetlands, on the other hand, appear to favor another undesirable invasive species, Phragmites australis (common reed grass).     

The lobed-lipped species of Haplochromis (Teleostei,Cichlidae) from Lake Edward,two instead of one

Thickened lips have evolved several times within the cichlid flocks of the East African Great Lakes. This distinct and easily recognisable phenotype is a model trait to study convergent evolution. Lake Edward (Eastern Africa) contains a unique cichlid assemblage, which has remained largely understudied. Hitherto, only one cichlid species with lobed lips, Haplochromis labiatus, was known from this lake. This species has a blunt snout and rounded and mostly retrognathous jaws. However, we found specimens with lobed lips, but with acute snouts and slender and mostly prognathous jaws. These belong to a species, hitherto unknown to science: H. lobatus sp. nov. Both species occur sympatrically. We evaluated the morphological diversity within these species by investigating a total of 112 specimens using traditional and geometric morphometric approaches. Both species are formally (re)described and show a relatively large variation in their trophic morphologies, which complicates identification. Both have a small gape, stout outer teeth that strongly decrease in size laterad, and a lower jaw set anteriorly with procumbently-implanted outer teeth. Haplochromis labiatus differs from H. lobatus sp. nov. by a straight to convex vs. straight to concave head, a slightly shorter and broader lower jaw (27.7–34.3 vs. 31.2–40.7% HL; 71.4–92.4 vs. 48.5–70.5% LJL), and lobed lips that are thickened uniformly over their whole lengths vs. medially enlarged. Gut content observations revealed that both species have an insectivorous diet. While both exploit similar food sources, their morphological differences presumably prevent them from entering into direct ecological competition.