Science and Research Information Report IR-16

Current and potential aquatic invasive species in Ontario and the Great Lakes region: A compilation of ecological information

Science and Research Information Report IR-16

Current and potential aquatic invasive species in Ontario and the Great Lakes region: A compilation of ecological information

Elizabeth C. Hatton1, Jeffrey D. Buckley1, Shannon A. Fera1,2, Samantha Henry1, Len M. Hunt3, D. Andrew R. Drake4 and Timothy B. Johnson1

1 Aquatic Research and Development Section, Ministry of Natural Resources and Forestry (MNRF), 41 Hatchery Lane, Picton, ON K0K 2T0 2 Current address: Fisheries Section, Species Conservation Policy Branch, MNRF, 300 Water Street, Peterborough, ON K9J 8M5 3 Centre for Northern Forest Ecosystem Research, MNRF, 103-421 James St S, Thunder Bay, ON P7E 2V6 4 Great Lakes Laboratory for Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, 867 Lakeshore Road, Burlington, ON L7S 1A1

2019

Science and Research Branch

Ministry of Natural Resources and Forestry © 2019, Queen’s Printer for Ontario

Copies of this publication are available from [email protected].

Cette publication hautement spécialisée, Current and Potential Aquatic Invasive Species in Ontario and the Great Lakes Region: A Compilation of Ecological Information, n’est disponible qu’en anglais conformément au Règlement 671/92, selon lequel il n’est pas obligatoire de la traduire en vertu de la Loi sur les services en français. Pour obtenir des renseignements en français, veuillez communiquer avec le ministère des Richesses naturelles et des Forêts au [email protected].

Some of the information in this document may not be compatible with assistive technologies. If you need any of the information in an alternate format, please contact [email protected].

Cite this report as: Hatton, E.C., J.D. Buckley, S.A. Fera, S. Henry, L.M. Hunt, D.A.R. Drake and T.B. Johnson. 2019. Current and potential aquatic invasive species in Ontario and the Great Lakes region: A compilation of ecological information. Ontario Ministry of Natural Resources and Forestry, Science and Research Branch, Peterborough, ON. Science and Research Information Report IR-16. 23 p. + appendices. Abstract

We reviewed agency watch lists, invasive species databases, published reports, and primary literature to identify and compile information on 73 fishes, 66 invertebrates, and 67 plants that have been identified as current or potential aquatic invasive species (AIS) to the Laurentian Great Lakes region. Nearly three quarters of the species are indigenous to Eurasia and are primarily associated with aquarium/water garden (61%) or commercial shipping (37%) pathways. Once established in the Great Lakes region, secondary spread is linked to recreational boating, canals, commercial shipping, and bait release. Body size ranges from microns to metres, although the majority are small or produce small seeds making detection and control challenging. Collectively, the species show a wide range of tolerances for temperature and salinity and will distribute across all types of aquatic habitat. Longevity ranges from days (invertebrates) to years (fish), with age at first maturity similarly ranging from ≈daily to several years. Plants are largely perennials (84%), are distributed among multiple growth habits, and most have both sexual (flowering) and asexual (predominantly fragmentation) reproductive strategies. Invertebrates reproduce sexually (98%), although some (24%) can also generate offspring asexually. Most fish species (56%) spawn without guarding their clutch, although many (36%) guard their young. Fish and invertebrates are predominantly omnivores, and some fish are carnivores, suggesting their trophic impact may be broad. Current and anticipated impacts of AIS include moderate to high competition for resources, nuisance growth (clogging infrastructure and waterways), declines in indigenous species, ecosystem change (including food webs and physical habitat), and new vectors for disease.

Résumé

Espèces aquatiques envahissantes actuelles et potentielles en Ontario et dans la région des Grands Lacs : une compilation de données écologiques

Nous avons passé en revue les listes de surveillance d’organismes, les bases de données sur les espèces envahissantes, les rapports publiés et la littérature (principaux travaux) afin de repérer et de compiler l’information sur 73 poissons, 66 invertébrés et 67 végétaux qui ont été identifiés comme étant des espèces aquatiques envahissantes (EAE) actuelles ou potentielles dans la région laurentienne des Grands Lacs. Près des trois quarts de ces espèces sont indigènes en Eurasie. L’invasion se fait surtout par des voies telles que les aquariums et les jardins aquatiques (61 %) ou la navigation commerciale (37 %). Une fois établie dans la région des Grands Lacs, la propagation secondaire est associée à la navigation de plaisance

Science and Research Information Report IR-16 iii et commerciale, aux canaux et au lâcher d’appâts vivants. La taille corporelle des espèces varie, de micromètres à mètres, bien que la plupart soient petites ou produisent de petites semences qui rendent difficiles la détection et le contrôle. Dans l’ensemble, les espèces montrent de vastes plages de tolérance aux écarts de température et de salinité, et elles se répandent dans tous les types d’habitats aquatiques. La longévité varie de jours (invertébrés) à années (poissons), et il en est de même pour l’âge à la première maturité, allant de ≈quotidienne (jour même) à plusieurs années. Les végétaux sont largement des vivaces (84 %) et se répartissent entre plusieurs types de croissance. La plupart ont des stratégies de reproduction à la fois sexuelles (la floraison) et asexuelles (surtout la fragmentation). Les invertébrés se reproduisent sexuellement (98 %), bien que certaines espèces (24 %) puissent aussi générer des progénitures de façon asexuelle. La plupart des espèces de poissons (56 %) se reproduisent sans garder leur couvée, mais bon nombre d’entre elles (36 %) veillent sur leurs petits. Les poissons et les invertébrés sont majoritairement omnivores, et certains poissons sont carnivores, ce qui indique un impact trophique susceptible d’être vaste. Les répercussions actuelles et anticipées des EAE comprennent une concurrence de modérée à élevée pour les ressources, une prolifération nuisible (engorgeant l’infrastructure et les voies navigables), le déclin d’espèces indigènes, des changements dans les écosystèmes (y compris les réseaux alimentaires et les habitats physiques) et de nouveaux vecteurs de maladies.

La base de données est disponible sur https://www.ontario.ca/fr/donnees/especes- aquatiques-envahissantes.

Acknowledgements

Funding for this study was provided by Great Lakes Protection Funding, which supports projects and activities that deliver on the commitments of the Ontario Ministry of Natural Resources and Forestry (MNRF) under the Canada-Ontario Agreement on Great Lakes Water Quality and Ecosystem Health and Ontario’s Great Lakes Strategy, as well as the Natural Heritage Policy Section of MNRF. Erika Jensen (Great Lakes Commission) shared information regarding U.S. species watch lists and contributed to preliminary discussions about this project. Allison Bannister (MNRF) contributed to a previous draft of this report, while Francine MacDonald (MNRF), Jeff Brinsmead (MNRF) and Shelley Arnott (Queen’s University) provided helpful suggestions on earlier drafts of this report.

Science and Research Information Report IR-16 iv Contents Abstract ...... iii Résumé ...... iii Acknowledgements ...... iv Background ...... 1 Methods ...... 3 Species selection ...... 3 Data collection ...... 5 Results ...... 7 Geographic ranges and pathways ...... 7 Species ecology ...... 9 Impacts ...... 14 Summary ...... 16 References ...... 19 Appendices map ...... 24 Appendix A. Methods ...... 25 Appendix B. Current and potential invasive fishes ...... 35 Appendix C. Current and potential invasive aquatic invertebrates ...... 78 Appendix D. Current and potential invasive aquatic plants ...... 117 Appendices references...... 158

Science and Research Information Report IR-16 v Science and Research Information Report IR-16 vi Background

The Laurentian Great Lakes Basin is one of the most heavily invaded freshwater ecosystems in the world with over 180 aquatic invasive species (AIS) reported since the mid-1800s (Ricciardi 2006, GLANSIS 2018). These species have global origins (Mills et al. 1993, Sturtevant et al. 2014, Davidson et al. 2017), with the rate of species discovery peaking in the early 1990s at 2.8 species per year (Mills et al. 2003, Ricciardi 2006, GLANSIS 2018). Species have arrived through a variety of pathways (e.g., commercial shipping, horticulture, pet and live food trades, canals and hydrological connections, recreational boating, authorized and unauthorized releases), with ballast water noted for the introduction of most AIS following the opening of the St Lawrence Seaway in 1959 (Mandrak and Cudmore 2010). While adoption of mandatory ballast water exchange rules in 2006 greatly reduced the risk associated with this pathway, many other pathways remain that can introduce and spread AIS (Ricciardi 2006, Bailey et al. 2011). The ecological impacts of AIS in the Laurentian Great Lakes have been substantial, and it is likely that the multitude of AIS has increased the scope and scale of ecological consequences (Ricciardi 2001) as evidenced by on-going food-web restructuring and other shifts in aquatic communities (Bunnell et al. 2014). The economic impact of AIS in the Great Lakes region (GLR) is also significant, estimated at over US$100 million annually (Rosaen et al. 2012).

Because established AIS can be nearly impossible to eradicate (Hobbs and Humphries 1995, Mack and Lonsdale 2002, Simberloff 2009) and the resulting costs of control can be substantial (Leung et al. 2002, Simberloff 2003, Rosaen et al. 2012), the most effective strategy to avoid the ecological and economic impacts of AIS is to prevent their arrival (Lodge et al. 2006). Research aimed at prevention has the immediate benefit of protecting the GLR from eventual ecological impacts, but also benefits the surrounding region given that the Great Lakes serve as a source for secondary invasions throughout North America, including the inland waters of Ontario (MacIsaac et al. 2004, Leung et al. 2006, Rothlisberger and Lodge 2013). Significant research has been undertaken to prevent the arrival of AIS, including identifying potential invaders to the GLR (Snyder et al. 2014, Pagnucco et al. 2015, Fusaro et al. 2016; see Table A1 for additional sources). However, as human populations, global trade, and recreational activity continue to expand, the risk of additional species accessing the GLR will increase. Despite the apparent success in greatly reducing the number of species arriving through ballast water (Briski et al. 2013), existing pathways for introduction and spread (e.g., recreational boating, live trade of aquatic foods and pets) lack similar management controls, while new pathways (e.g., internet trade of live animals; Padilla and Williams 2004) may shift the future arrival and spread of key species. The arrival of species to the GLR will also change due to shifting human and environmental conditions. For

Science and Research Information Report IR-16 1 example, new patterns of species introductions are expected because of changes to human population density and resulting changes to existing pathways for AIS in the region. Climate change will facilitate the survival of species in new regions, including by altering the suitability of aquatic habitats for AIS relative to indigenous taxa (Sharma et al. 2007, Walther et al. 2009, Lynch et al. 2010, Alofs et al. 2014), thereby shifting the intensity and spatial distribution of ecological effects.

Effective management of AIS requires that species likely to arrive, survive, establish, and impact the GLR be identified with reasonable certainty. Risk assessments that consider species life history traits, habitat suitability, historical patterns of invasion, and ecological impacts in other invaded regions have become critical resources to identify and prioritize species of concern (reviewed in Dahlstrom et al. 2011, Verbrugge et al. 2012). Numerous risk assessments have been developed for individual species (Mandrak et al. 2012, Cudmore et al. 2012), groups of species (Kolar and Lodge 2002, Gantz et al. 2015), and pathways (Bailey et al. 2011, Drake and Mandrak 2014, Drake et al. 2017), and have been used, in conjunction with other research and management exercises, to generate regulated and watch lists of species that pose a high likelihood of arriving and establishing in the GLR (see Table A1). More recently, efforts have sought to develop risk frameworks to predict AIS establishment and impacts across multiple taxonomic groups and pathways (Sturtevant et al. 2014, Nienhuis and Haxton 2016, Davidson et al. 2017) and to develop generalized species niche avatars (Larson et al. 2014). Such generalized risk assessment approaches may reduce dependence on detailed information (i.e., species-specific biology, habitat preferences, and physiological tolerances to be matched to species distributions and pathway movement), however, sufficient information for assessments may either be lacking (Fusaro et al. 2016) or collecting this information can take months or years (Lodge et al. 2016).

As part of a larger effort to develop a risk assessment tool to predict the distribution and spread of AIS in Ontario, we compiled information about current and potential AIS in Ontario and the GLR, focusing on their geographic origins, likely pathways of arrival and spread, ecological characteristics, and generalized ecological impacts. The review was based on databases, published literature, and readily accessible reports, and was developed to gain a better understanding of the variety of AIS likely to colonize Ontario and the GLR. We refer the reader to the original data sources for more in-depth understanding of individual species.

Science and Research Information Report IR-16 2 Methods

Species selection

We compiled a list of 206 non-indigenous fishes, aquatic invertebrates, and aquatic plants that have been identified as current or potential aquatic invasive species (AIS) in Ontario or the Great Lakes region (GLR) based on existing data sources. We defined the GLR to include the Great Lakes Basin, as well as all inland waters of adjoining provincial and state territories (i.e., Canada including Manitoba, Ontario, Quebec, and United States of America including Illinois, Indiana, Michigan, Minnesota, New York, Ohio, Pennsylvania, Vermont, Wisconsin).

The species list was developed in 2015 by summarizing species found on regulated lists (primarily government documents, including species prohibited for possession or import in a given jurisdiction) and watch lists (primarily non-government sources, designed to identify future species of concern), invasive species databases describing the current distribution of AIS in the region, databases of primary literature and risk assessments, as well as primary literature describing or predicting potential species invasions. The full list of data sources is presented in Table A1. Many species were identified by multiple sources; the basis of listing a species was recorded for primary or grey literature only when not superseded by a regulated or watch list. Noteworthy is that the methods used in each data source varied and do not necessarily represent a consistent approach to assessing a species’ invasion potential. For example, the methods used to assess species arrival, survival, establishment, spread, and ecological impact varied widely; in some cases, only a subset of these steps were used to identify species posing invasion risk to the region.

For those species already present in the GLR, we used year of establishment and status to identify species that should remain in the analysis. For example, species that have been observed but never established were not included unless a risk assessment indicated climate change may affect their potential to establish.

The Great Lakes Aquatic Nonindigenous Species Information System (GLANSIS; 2018) identified over 180 non-indigenous species established in the Great Lakes Basin. We included 57 of these species in our assessment because they have had substantial detrimental impacts (e.g., Dreissena polymorpha and D. bugensis (zebra and quagga mussels)), their populations are actively being controlled to minimize impacts (e.g., Petromyzon marinus (sea lamprey)), and they are still very likely to continue to spread (e.g., Neogobius melanostomus (round goby), Myriophyllum spicatum (Eurasian watermilfoil)).

Science and Research Information Report IR-16 3 The final species list was developed based on criteria involving 2 distinct categories of species (Figure 1):

1) Current AIS: aquatic invasive species for which evidence of established populations exists in Ontario or the GLR that could spread in the region

2) Potential AIS, not established in the GLR:

a) aquatic invasive species which are established elsewhere in North America, that may spread into the GLR through natural dispersal or human-mediated movement

b) aquatic invasive species for which none or little evidence of establishment exists in North America but have been identified as invasive elsewhere in the world, and whose thermal requirements align with the current climate of Ontario or the GLR or conditions projected under climate change.

Figure 1. Conceptual diagram reflecting the general categories of aquatic invasive species (AIS) used in this analysis, based on indigenous origin and pathways.

Where appropriate, species thermal habitat requirements were compared to the long- term (2071–2100) scenarios from the Intergovernmental Panel on Climate Change’s Fifth Assessment Report, which show air temperature increasing from 2.3 to 7.9 °C across the Great Lakes Basin. Species remained on the list if they matched projected thermal conditions, even if that habitat was not currently available in Ontario. Thermal matching criteria were based on species’ lower or upper thermal limits for survival or thermal guild (see below) and air temperature was used as a proxy for water temperature. In a similar manner, species whose thermal habitat was classified as

Science and Research Information Report IR-16 4 temperate (with a southern distribution) or sub-tropical were considered as possible AIS if they have shown high plasticity to a variety of thermal environments or were known to be highly invasive.

Given the constraints associated with the availability of data, we acknowledge that the list of species retained for analysis does not include all possible AIS to the region. While our list contains 206 species, approximately 1,000 freshwater and brackish aquatic species could establish in the GLR (CABI ISC 2018). To the best of our knowledge, the species retained in our analysis, or those very similar to these species, pose the greatest invasion risk now and into the future.

Data collection

Once the master list of species was identified, detailed information about each species was compiled from databases and reports in 2018 (listed in Table A2). These data included indigenous and invaded ranges, current jurisdictional prohibitions and restrictions (if any), pathways of initial introduction and secondary spread, ecological factors (including habitat, diet and reproduction), and ecological impacts. Rather than providing a detailed account for each species, we compiled summaries and present a synthesis for each species to summarize and standardize the data for comparison among taxa. The data fields compiled for this report are described in Table A3. A list of abbreviations used in this report is presented in Table A4. The species-specific data are summarized in appendices B (fishes), C (aquatic invertebrates), and D (aquatic plants). The data set (*.csv format) can be accessed at https://ontario.ca/data/aquatic-invasive- species.

For many fields, species had multiple entries (e.g., indigenous range included both Europe and North America, potential pathway of entry could be via commercial shipping or aquarium/garden trade) and in these cases entries were counted separately and were not ranked or presented in a priority sequence. When it came to classifying indigenous geographic ranges, Eurasia was occasionally reported in place of Asia or Europe; in these cases Eurasia was retained, as it was not possible to resolve if species’ origins were singular to Europe or Asia or indigenous to both.

The actual or potential primary pathway(s) of initial entry into the GLR, as well as potential or actual GLR spread pathways for each species were categorized as: • aquarium/garden (species popular in the aquarium/pet trade, ornamental water gardens, and plant trade) • bait release (including water in which the bait was held) • canal (dispersal by waterway connections to GLR) • commercial shipping (Great Lakes and transoceanic ballast water and fouling)

Science and Research Information Report IR-16 5 • escaped culture (unintentional commercial release) • hitchhiker (with other organisms or goods) • intentional release (authorized stocking) • live trade (including aquaculture, bait farms/retailers, fish markets/restaurants) • recreational boating (including live wells, motors, transom wells, bilges, trailers, and equipment) • unauthorized intentional release (including dumping of unwanted plants or animals, ceremonial release) • unknown

For many species, multiple pathways had been identified as probable and all were included in our analysis. Many of the invertebrate and plant species we analyzed have small adult or juvenile body sizes or the ability to produce resting stages or seeds, increasing their potential to enter and spread in the GLR as hitchhikers with other organisms or goods in trade. Additional pathways not considered in our study include movement of waterfowl, float planes, and commercial or agency research activity/equipment because they represent low frequency events.

Temperature tolerance was summarized only for fishes because temperature is an abiotic master factor for them (Brett 1971, Fry 1971) and their physiological performance is optimized across a narrow temperature range (Beitinger and Fitzpatrick 1979, Pörtner and Farrell 2008). Fishes were assigned thermal guilds based on Hatton et al.’s (2018) derived final temperature preferendum (FTP) and Coker et al.’s (2001) classification: warm (FTP >25 °C), cool (FTP 19–25 °C), and cold (FTP <19 °C). Two intermediate classes (i.e., warm/cool or cool/cold) were selected for fish with overlapping FTP values.

To better understand potential natural dispersal of fishes across the landscape, fish species were assigned migratory classifications (e.g., anadromous (migrates from salt to fresh water), potamodromous (migrates in fresh water only)). However, some species have the physiological capacity/plasticity to adapt and change migratory behaviour based on exposure to different habitat conditions (e.g., landlocked freshwater versus anadromous alewife, Alosa pseudoharengus; Palkovacs et al. 2008). Fish species were also grouped by reproductive guild based on reproductive behaviour (as noted in Coker et al. 2001, Froese and Pauly 2017, IUCN 2018) and Balon’s (1975, 1981) classification of spawning behaviour.

Science and Research Information Report IR-16 6 Results

The final list of AIS contained 206 species: 73 fishes, 66 invertebrates, and 67 plants (see appendix tables B1, C1, and D1 for fishes, invertebrate, and plant species lists, respectively). Of these 206 species, 41% were established (currently present) in one or more Great Lakes provinces or states (Table 1). Over half (57%) of the species we analyzed were regulated (prohibited or restricted use) in at least one province or state. Of the AIS not established, 33, 13, and 100% of fish, invertebrate, and plant species, respectively, are regulated in at least one province or state. Some snakehead and Asian carp species are prohibited or regulated in all jurisdictions.

Table 1. Aquatic invasive species (AIS) count for Ontario and the Great Lakes region (GLR) by current (2018) distribution category. Distribution Fishes Invertebrates Plants Total count Current AIS: present in Ontario or other Great Lakes region (GLR) 28 21 35 84 jurisdiction(s) Potential AIS: not established in GLR, 17 13 25 55 present in North America Potential AIS: not established in North 28 32 7 67 America, invasive elsewhere Total count 73 66 67 206

Geographic ranges and pathways

Species included in our analysis had indigenous geographic ranges across six continents, with most species (75%) having ranges in Eurasia (including Europe and Asia). Eurasian species made up 77% of the species that are currently established and 73% of potential AIS in the GLR (Figure 2). Thirty-eight species (19%) were indigenous to North America and included several species native to one or more Great Lakes provinces or states, but not all (e.g., Fundulus diaphanus (banded killifish), Cabomba caroliniana (fanwort)). Figure 3 shows the indigenous range of all species and the movement and establishment on other continents; species can originate from their indigenous range(s), as well as non-indigenous established range(s).

Initial pathways of entry into the GLR for potential invasive species were primarily associated with aquarium/garden, commercial shipping, and live food (Figure 4). Once established, movement and spread of both current and potential invasive species are linked to recreational boating, bait release, canal, and commercial shipping pathways.

Science and Research Information Report IR-16 7 Compiled species-specific information on ranges and pathways is provided in tables B2, C2, and D2.

Figure 2. Indigenous geographic ranges of current (2018) or potential aquatic invasive fish, invertebrate, and plant species (AIS) in Ontario and the Great Lakes region.

Figure 3. Source of current (2018) and potential aquatic invasive fish, invertebrate, and plant species (AIS) to Ontario and the Great Lakes region (GLR) by indigenous continent. Circle and arrow colour signify the continent of origin, circle size represents the number of current or potential AIS originating from that continent, and arrow width and direction reflect the number of species that have, or are likely to, move from that continent to Ontario and the GLR.

Science and Research Information Report IR-16 8 Figure 4. Initial pathways of entry (primary introductions) to the Great Lakes region (GLR) and secondary spread (movement within the GLR) for current (2018) and potential aquatic invasive fish, invertebrate, and plant species (AIS).

Species ecology

Across all species, physical size was highly variable (Figure 5). Maximum reported adult sizes (Table 2) range from species found in a drop of water (Prymnesium parvum (golden alga)) to very large fish (Silurus glanis (Wels catfish)). While information on maximum adult length was readily reported for most species, records of maximum weight were less available. Fish body weight was most frequently reported (41% of the time; Table B3).

Numerous aquatic habitats (Figure 6) were identified as having the potential to support AIS, including rivers/streams, lakes/ponds, wetlands, and riparian zones (plants only). Detailed descriptions of habitat preference are provided in the appendices (Tables B3- D3). Growth habits of the plants included in our analysis varied from submergent to

Science and Research Information Report IR-16 9 riparian (Table 3), with 52% of species able to develop more than one growth form (e.g., Crassula helmsii, Australian stonecrop).

Figure 5. Maximum adult size (cm, length or height) for current (2018) and potential (combined) fish, aquatic invertebrate, and aquatic plant species that are non-indigenous to the Great Lakes region. The box plot represents the median (middle line), interquartile range (box), first and third quartiles (the 25th and 75th percentiles; lower and upper hinges), smallest and largest points within 1.5x the interquartile range from the hinges (whiskers), and outliers (more than 1.5x the interquartile range; dots).

Table 2. Adult size ranges by taxonomic group for current (2018) and potential (combined) aquatic invasive fish, invertebrate, and plant species in the Great Lakes region. Adequate weight data were only available for fishes. n = number of reported values for each taxonomic group Species Minimum adult size Maximum adult size group Hyrcanogobius bergi Silurus glanis 1.6 cm 5 m Fishes (Volga dwarf goby) (Wels catfish) (n=73) Alburnus alburnus S. glanis 60 g 306 kg (common bleak) (Wels catfish) Invertebrates Filinia spp. Schyzocotyle acheilognathi <200 µm 1 m (n=66) (wheel animal) (Asian fish tapeworm) Plants Prymnesium parvum Cabomba caroliniana 10 µm 10 m (n=64) (golden alga) (fanwort)

Science and Research Information Report IR-16 10 Figure 6. Habitat preference of current (2018) and potential aquatic invasive (AIS) fish, invertebrate, and plant species in the Great Lakes region.

Table 3. Plant growth habits of current (2018) and potentially invasive aquatic species (AIS) in the Great Lakes region.

Growth habit Current AIS Potential AIS Free floating 6 10 Submergent 14 10 Emergent 17 23 Floating leaf 10 16 Riparian 13 8

The species varied greatly in their tolerances to salinity and water temperature. Many of the fish, invertebrate, and plant species tolerate a wide range of salinities (Figure 7). For fish thermal preferences, distribution across thermal guilds was even, with warm, cool, and cold representing 33, 29, and 32% of the fishes, respectively (Table B3).

Omnivory was the most common trophic guild for the fish and invertebrate species included in our analysis (Figure 8). Fishes were most commonly classified as omnivores or carnivores. For invertebrates, 58% were omnivorous and 24% herbivorous, with the remainder made up of a few carnivores and parasites. More in depth description of each species’ trophic guild and diet is provided in tables B4, C4, and D4. All plants were identified as autotrophs, except for Pfiesteria spp. (phantom dinoflagellate) which can also function as a heterotrophic omnivore.

Science and Research Information Report IR-16 11 Figure 7. Salinity tolerance of current (2018) and potential aquatic invasive (AIS) fish, invertebrate, and plant species in the Great Lakes region. F, B, and M indicate freshwater, brackish, and marine habitat, respectively.

Figure 8. Trophic guilds of current (2018) and potentially invasive aquatic fish and invertebrate species (AIS) in the Great Lakes region.

Science and Research Information Report IR-16 12 Age at reproductive maturity and longevity for fish and invertebrates was highly variable, spanning days to decades (Table 4). Mode of reproduction varied among taxonomic groups. All fishes reproduced sexually, although gibel carp (Carassius gibelio) can also reproduce asexually. Examining reproductive guilds for fish, 56% of fish spawned without guarding their clutch, the remaining either guarding (36%) or bearing (8%) their young live. Sexual reproduction was common for the invertebrates (97%), while asexual reproduction was possible for some species (24%). For plants, most species (85%) could reproduce both sexually and asexually and most were classified as perennials (84%). Asexual reproductive strategies for plants included vegetative fragmentation and offsets from rhizomes and stolons (Table 5). Compiled species-specific reproduction information can be found in tables B4, C4, and D4.

Table 4. Examples of minimum and maximum age at reproductive maturity and longevity of current and potential invasive fishes and aquatic invertebrates to the Great Lakes region. n=number of reported values for each taxonomic group.

Fishes Invertebrates (n=62) (n=56) 1–3 months 1 day Min Gambusia affinis Filinia spp. (western mosquitofish) (wheel animal) Age at maturity 5–6 years 4 years Max Hypophthalmichthys harmandi Astacus astacus (largescale silver carp) (European crayfish) 1 year 3–15 days Min Hyrcanogobius bergi Cercopagis pengoi (Volga dwarf goby) (fishhook water flea)

Longevity 13–38 years 20 years Max Cyprinus carpio A. astacus (common carp) (European crayfish)

Science and Research Information Report IR-16 13 Table 5. Reproductive strategies of current and potential aquatic invasive plant species (AIS) in the Great Lakes region. Reproductive Current AIS Potential AIS Reproduction strategy count count Sexual External pollination 28 19 Self-pollination 7 7 Asexual Fragmentation 34 31 Rhizomes 17 9 Stolons 9 11 Turions 4 2 Tubers 1 3 Bulbs 1 0

Impacts

Resource competition with indigenous species (either in North America or other invaded regions) was the most commonly reported ecological impact across all species but most species had multiple impacts (Figure 9). Impacts of forty-five species were unknown or unidentified primarily due to inadequate species-specific research.

Of the 206 species of concern, GLANSIS technical species profiles were available for 118; the assessed impacts of these species are summarized in Figure 10. Most of the environmental effects documented for these species were negative. For most of the socio-economic impacts and beneficial outcomes (including advantageous effects on the environment or humans), little to no evidence was available to support significant impacts, indicating a lack of information and understanding of potential socio-economic impacts and beneficial effects of these species. Species-specific impacts are compiled in tables B4, C4 and D4.

Science and Research Information Report IR-16 14

Figure 9. Summary of potential impacts of current (2018) and potential aquatic invasive fish, invertebrate, and plant species (AIS) in the Great Lakes region as gathered from databases and reports (see Table A2).

Figure 10. Ranked impact assessment of current (2018) and potential aquatic invasive fish, invertebrate, and plant species (AIS) in the Great Lakes region (as qualified by GLANSIS).

Science and Research Information Report IR-16 15

Summary

We consolidated and classified characteristics of aquatic species that have been identified as current or potential AIS to the GLR. Many online and published resources provide detailed accounts for individual species (see Appendix A1). However, we used standardized language for common factors among species to help identify patterns within and among taxonomic groups.

Of the 206 species listed (as of December 2015), 41% were already established in at least one Great Lakes province or state and may pose invasion risk to other jurisdictions. Only 57% of the listed species are either prohibited or regulated for specific use (e.g., live food trade) in at least one jurisdiction. Most species are native to Eurasia, with commercial shipping, the aquarium and garden trade, and live food trade as the most probable pathways for primary introduction. Commercial shipping is the pathway of entry primarily associated with listed invertebrates while the initial introduction of fishes and plants is associated with the pet, aquarium, and live food trades. Once established, secondary spread throughout the GLR for all species is via recreational boating, canals, and commercial shipping.

Across taxonomic groups, all aquatic habitats are vulnerable to invasion. Most listed plants are emergent perennials, although submergent, floating, and riparian growth forms are also represented. Most fish are carnivorous or omnivorous suggesting the potential for competition with many socially and economically valuable native fishes. Most invertebrates are omnivores so the likely impact is resource competition with native species. Most of the plants are expected to compete with native plants or alter their habitats, which will have broader ecosystem consequences. Efforts have mainly been directed at understanding possible negative environmental impacts if the species arrived and established, with limited research on socio-economic impacts or potential beneficial outcomes (e.g., commercial and sport fisheries, prey/plants supporting fisheries).

Once a species arrives in the GLR, managing the risk of establishment and spread is critical. Very few control mechanisms exist and, when available, are rarely species- specific; they typically are generic chemical controls (e.g., broad-spectrum herbicides such as diquat) and insecticides/piscicides (e.g., rotenone). Widespread application of these management tools is restricted by strict regulation in most jurisdictions, combined with high expense due to the need for repeated applications, concerns about non-target or downstream environmental effects, and the potential for species resistance. Biological, physical, and mechanical controls have been implemented, sometimes with containment success but evidence of effectiveness is lacking (Rytwinski et al. 2018), they may carry unintended or undesirable effects (Kopf et al. 2017), and the lag

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between arrival and early detection in large dynamic ecosystems such as the Great Lakes often makes eradication unattainable.

Risk assessments are necessary to predict the arrival, survival, establishment, spread, and impacts of AIS. Species-specific risk assessments have been developed for about half of the species in our analysis (75, 32, and 42% of fish, invertebrate, and plant species, respectively). Typically, risk assessments for AIS incorporate the processes of species arrival, survival, establishment, and spread, as well as the magnitude of ecological consequences should these processes occur (MNRF guidelines, Nienhuis and Haxton 2016; DFO guidelines, Mandrak et al. 2012). However, the scope of and detail in risk assessments varies. In addition, methods varied among the studies included in our analysis, which affects how the results can be interpreted. For example, some risk assessments omit the process of species arrival, which results in a framework to prioritize species based on the potential for survival and establishment, but not necessarily based on whether species are likely to arrive or even be present in a transport pathway. Such approaches are appropriate to develop import regulations by identifying species with the potential to survive and establish. However, they may not be suited for prioritizing surveillance activities in a recipient region, given that a species may have a low likelihood of arrival. In other cases, ecological effects may not have been explicitly incorporated in risk estimates. Meaningful comparison of risk assessment results across species requires understanding and accounting for the differences in methods. In this report, we incorporate all risk assessment results and associated species listing processes, but caution that risk rating comparisons across species requires consideration of the risk assessment methods involved.

With over 180 non-indigenous species already present in the GLR, at an estimated US$100 million annual cost to manage and compensate the associated harm (Rosaen et al. 2012), prevention of species introductions remains the most economically effective strategy (Lodge et al. 2006). Advances in modern risk assessment tools, including bio- economic analysis to better understand trade-offs between prevention and control (Leung et al. 2002), will provide decision makers with a better understanding of the consequences of alternative management strategies (Lodge et al. 2016). Continued research to improve forecasting models, combined with new and more cost-effective surveillance tools will allow management to focus surveillance, outreach, and control efforts on the highest risk species and locations. Ongoing ecological change, including climate change, population growth, changing commerce (e.g., growth in aquaculture, exotic pet trade), and continued globalization will only amplify the potential for non- indigenous species to be introduced, but the economic and ecological harm can be reduced or avoided through continued education and prevention initiatives.

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Synopsis:

• 73% of potential AIS to Ontario and the GLR have an indigenous range that includes Eurasia.

• Initial pathways of introduction are primarily associated with the aquarium/garden trade, commercial shipping, and live food.

• Once an AIS establishes in the GLR, pathways of continued spread are linked to recreational boating, canals, commercial shipping, and bait release.

• Many current and potential AIS are of small body size (<1 cm) or produce juvieniles, small seeds or resting stages that are difficult to detect and can be readily transported via many pathways.

• Many current and potential AIS tolerate a range of habitats, salinities, and prey/food items.

• Greatest AIS knowledge gaps exist for invertebrate and plant species, while potential fish species are generally better understood.

• Current and anticipated adverse ecological impacts of AIS include moderate to high resource competition, nuisance growth (clogging of infrastructure and waterways), indigenous species decline, ecosystem change, and new vectors for disease.

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Appendices map

Appendix Table Section Contents Page number

A 1 Methods Species listing references and decisions 26

2 Search framework/references 28

3 Database field definitions 31

4 Common acronyms 34

B 1 Fishes Taxonomy 35

2 Range and pathways 38

3 Size and habitat 51

4 Diet and reproduction 60

5 Impact 67

C 1 Invertebrates Taxonomy 78

2 Range and pathways 82

3 Size and habitat 93

4 Diet and reproduction 100

5 Impact 105

D 1 Plants Taxonomy 117

2 Range and pathways 120

3 Size and habitat 133

4 Growth habit and reproduction 139

5 Impact 144

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Appendix A. Methods

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Table A1. References used to identify aquatic invasive species in Ontario and the Great Lakes region, listed alphabetically. n/a=not applicable

Jurisdiction/organization Citation/website Regulated and Watch Lists: California State Noxious Weed List https://plants.usda.gov/java/noxious?rptType=State&statefips=06 https://www.cityofchicago.org/dam/city/depts/doe/general/NaturalResourcesAndWaterC Chicago (shipping canal concern list) onservation_PDFs/InvasiveSpecies/InvasiveSpeciesRulesAndRegulations52011.pdf European Union list http://ec.europa.eu/environment/nature/invasivealien/list/index_en.htm https://www.flrules.org/gateway/readFile.asp?sid=0&tid=5973277&type=1&File=5B- Florida 64.011.doc GLANSIS watch list https://www.glerl.noaa.gov/glansis/nisListGen.php Great Lakes-St. Lawrence Governor’s and http://www.gsgp.org/media/2017/ais-least-wanted-announcement.pdf Premiers “Least Wanted” List Illinois https://www.invasive.org/species/list.cfm?id=152 Indiana https://www.in.gov/dnr/3123.htm Michigan https://www.michigan.gov/invasives/0,5664,7-324-68002_74188---,00.html https://whttps://www.michigan.gov/invasives/0,5664,7-324-68002_74188--- Minnesota ,00.htmlww.dnr.state.mn.us/invasives/laws.html#prohibited New York http://www.dec.ny.gov/docs/lands_forests_pdf/islist.pdf Ohio http://ohiodnr.gov/ais https://www.ontarioinvasiveplants.ca/wp-content/uploads/2016/07/2016-Aquatic-Watch- Ontario Invasive Plants Council watch list List_-Final.pdf Ontario prohibited and restricted species list https://www.ontario.ca/page/stop-spread-invasive-species (shortlist) Texas Invasive Species Institute http://www.tsusinvasives.org/home/

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Jurisdiction/organization Citation/website US Department of Agriculture list https://www.invasivespeciesinfo.gov/aquatics/main.shtml US Fish and Wildlife Service screening list https://www.fws.gov/fisheries/ANS/erss_high_risk.html Wisconsin https://dnr.wi.gov/topic/Invasives/documents/nr40lists.pdf Invasive species databases and databases of primary literature and risk assessments: CABI ISC https://www.cabi.org/isc/ EDDMapS http://www.eddmaps.org/ontario/distribution/ GLANSIS https://www.glerl.noaa.gov/glansis/ Global Invasive Species Database http://www.iucngisd.org/gisd/ USGS NAS https://nas.er.usgs.gov/ Primary literature: n/a Marcogliese et al. 2016 n/a Pagnucco et al. 2015 n/a Aguilera 2015 n/a Straka et al. 2015 n/a Snyder et al. 2014 n/a Herborg et al. 2007 n/a Kolar and Lodge 2002

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Table A2. Databases and reports used in species-specific data syntheses (accessed March through October 2018) of aquatic invasive species in Ontario and the Great Lakes region, listed alphabetically. n/a=not available.

Database or Citation Website accessed report APIRS. 2018. Aquatic and Invasive Plant Information Retrieval System. APIRS https://plants.ifas.ufl.edu/apirs/ University of Florida, Gainesville, FL. AquaNIS, Editorial Board. 2015. Information system on Aquatic Non- http://www.corpi.ku.lt/databases/index.php/aqua AquaNIS Indigenous and Cryptogenic Species. World Wide Web electronic nis publication. Version 2.36+. CABI ISC. 2018. Invasive Species Compendium. CAB International. CABI ISC https://www.cabi.org/isc/ Wallingford, UK. DAISIE. 2018. Delivering Alien Species Inventories for Europe species DAISIE http://www.europe-aliens.org factsheets. DORIS. 2018. Données d'Observations pour la Reconnaissance et DORIS http://doris.ffessm.fr l'Identification de la faune et la flore Subaquatiques. Froese, R. and D. Pauly, editors. 2018. Fishbase, version 10. World Wide FishBase http://www.fishbase.ca/ Web electronic publication. GISD GISD. 2018. Global Invasive Species Database. http://www.iucngisd.org/gisd/ GLANSIS, 2018. Great Lakes Aquatic Nonindigenous Species Information System. US Geological Survey, Nonindigenous Aquatic Species Database, https://www.glerl.noaa.gov/glansis/nisListGen.ph GLANSIS Gainesville, FL, and NOAA Great Lakes Aquatic Nonindigenous Species p Information System, Ann Arbor, MI. Invasive Plant Swearingen, J. and C. Bargeron. 2016 Invasive Plant Atlas of the United Atlas of the States. University of Georgia Center for Invasive Species and Ecosystem https://www.invasiveplantatlas.org United States Health.

ITIS ITIS. 2018. Integrated Taxonomic Information System (ITIS). http://www.itis.gov

IUCN. 2017. International Union for Conservation of Nature Red List of IUCN Red List http://www.iucnredlist.org/ Threatened Species. Version 2017-3.

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Database or Citation Website accessed report NAS. 2018. Nonindigenous Aquatic Species Database. US Geological NAS/USGS https://nas.er.usgs.gov Survey, Gainesville, FL. Fofonoff P.W., Ruiz G.M., Steves B., Simkanin C., and J.T. Carlton JT. NEMESIS 2018. National Exotic Marine and Estuarine Species Information System https://invasions.si.edu/nemesis/ (NEMESIS). NISIC. 2018. National Invasive Species Information Center. National NISIC https://www.invasivespeciesinfo.gov/aquatics/ Agricultural Library, US Department of Agriculture, Beltsville, MD.

NOBANIS NOBANIS. 2018. European Network on Invasive Alien Species factsheets. https://www.nobanis.org/

OISAP. 2018. Ontario Invading Species Awareness Program. Ontario OISAP Federation of Anglers and Hunters, Ontario Ministry of Natural Resources http://www.invadingspecies.com/ and Forestry. Rotifer World Jersabek, C.D. and M.F. Leitner. 2018. The Rotifer World Catalog. World http://www.rotifera.hausdernatur.at Catalog Wide Web electronic publication. TISI TISI. 2014. Texas Invasive Species Institute. http://www.tsusinvasives.org USFWS. 2018. US Fish and Wildlife Service, Ecological Risk Screening https://www.fws.gov/fisheries/ans/species_erss_ USFWS Summary Reports. reports.html Courtenay, W.R. and J.D. Williams. 2004. Snakeheads (Pisces, USGS Channidae): A biological synopsis and risk assessment. US Geological n/a Survey, Geological Survey Circular 1251. 143 p. Kolar, C.S., D.C. Chapman, W.R. Courtenay Jr., C.M. Housel, J.D. Williams and D.P. Jennings. 2005. Asian carps of the genus https://digitalcommons.unl.edu/cgi/viewcontent.c USGS Hypophthalmichthys (Pisces, Cyprinidae) ― A biological synopsis and gi?article=1004&context=natlinvasive environmental risk assessment. US Fish and Wildlife Service. 185 p. Free-living and Parasitic Copepods (Including Branchiurans) of the https://www.glsc.usgs.gov/greatlakescopepods/ USGS Laurentian Great Lakes: Keys and details on individual species MainMenu.php WIDNR. 2018. Wisconsin Department of Natural Resources — Aquatic Wisconsin DNR https://dnr.wi.gov/topic/Invasives/ Invasive Species Literature Reviews. WoRMS WoRMS Editorial Board. 2018. World Register of Marine Species. http://www.marinespecies.org

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Table A3. Description of database fields used to summarize distribution, ecology, and impacts of current and potential aquatic invasive species in the Great Lakes region. n/a=not applicable.

Field name Description Filtered response(s) Taxonomy Scientific name n/a n/a Common name(s) Names most commonly associated with species n/a Family/Organism Category for classifying or grouping taxa (e.g., n/a category taxonomic family or plant type) ITIS TSN ITIS taxonomic serial number n/a Reason for inclusion, including proposed regulations, restrictions, watch lists, possession Basis for listing n/a bans, present in GLR, present in North America, scientific literature Origin and range Indigenous geographic range of species by Africa, Asia, Eurasia, Europe, Oceania, North America, South Origin continent America Indigenous geographic range by area or country, Origin detail n/a when found GLR regulated GLR jurisdictions where species is regulated Great Lakes region provinces and states (ON, PQ, IL, IN, MI, locations (prohibited or restricted use) NY, OH, PA, VT, WI) Year species first observed in GLR, established GLR year n/a populations only Great Lakes (list), Great Lakes region (ON, PQ, IL, IN, MI, Current GLR range Location(s) where species is found in GLR NY, OH, PA, VT, WI) Non-indigenous geographic range by continent Africa, Asia, Eurasia, Europe, Oceania, North America, South Current invaded range (current global distribution = origin + current America range) aquarium/garden, bait release, canal, commercial shipping, Actual or potential initial entry pathway(s) to the escaped culture, hitchhiker, intentional release, live trade, Introduction pathway GLR; does not include secondary spread recreational boating, unauthorized intentional release, unknown

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Field name Description Filtered response(s) aquarium/garden, bait release, canal, commercial shipping, Actual or potential secondary spread pathway(s) escaped culture, hitchhiker, intentional release, live trade, GLR spread pathway in the GLR, once species has established recreational boating, unauthorized intentional release, (includes original initial entry pathways) unknown Species ecology Typical length of largest/oldest adult of the Maximum adult length n/a species; units specified for each record Typical weight of largest/oldest adult of the Maximum adult weight n/a species; units specified for each record Typical maximum lifespan; units specified for Longevity n/a each record Preferred temperature for fishes (Hatton et al. Thermal guild cold, cold/cool, cool, cool/warm, warm 2018) Salinity tolerance General classification of salinity tolerated freshwater, brackish, marine Type of habitat typically occupied for most of the Habitat preference river/stream, lake/pond, wetland, riparian year pelagic, benthic, littoral, benthopelagic, parasitic; a variety of Habitat description Spatial position in habitat descriptors for soil/substrate conditions for plants Trophic guild Primary mode of feeding, position in food web omnivore, herbivore, carnivore, parasite, unknown plankton, phytoplankton, zooplankton, detritus, benthic Diet details Common prey identified in diet macroinvertebrates, fish, small fish, fish eggs, algae, macrophytes, insects, amphibians Typical length of time to reproductive age; units Age to maturity n/a specified for each record Mode of reproduction, including migratory sexual, asexual; amphidromous, anadromous, catadromous, Reproduction behaviour where applicable non-migratory, potamodromous General strategies for plant reproduction, external pollination, self-pollination; bulbs, rhizomes, stolons, Reproduction strategy including both sexual and asexual methods tubers, turions, fragmentation nonguarder, broadcast spawner; nonguarder, brood hider; Reproductive behaviour and habitat for fishes Reproduction guild guarder, substrate chooser; guarder, nest spawner; bearer, (Balon 1975) external; bearer, internal

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Field name Description Filtered response(s) General classification of plant species growth emergent, submergent, floating, floating leaf, free floating, Growth habit forms riparian, epiphytic Life cycle General classification of plant species life cycles annual, perennial Impacts Negative impacts, ranked assessment (source: high, moderate, little/no evidence, inadequate research, Environmental impact GLANSIS impact assessments) unavailable Socio-economic Negative impacts, ranked assessment (source: high, moderate, little/no evidence, inadequate research, impact GLANSIS impact assessments) unavailable Positive effects, ranked assessment (source: high, moderate, little/no evidence, inadequate research, Beneficial effects GLANSIS impact assessments) unavailable competition, predation, ecosystem change, habitat alteration, native species declines, disease vector, increase Impact summary Summary of actual or potential impacts contaminant flow, hybridization risk with indigenous species, nuisance species, toxic, not likely, unknown Impacts and possible Specific impact details (actual and potential) n/a interactions Listed agencies or jurisdictions with available Risk assessment n/a risk assessments

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Table A4. Abbreviations that appear in this report.

Abbreviation Definition AIS aquatic invasive species B brackish (salinity) CA California Centre for Agriculture and Bioscience International CABI ISC Invasive Species Compendium CFIA Canadian Food Inspection Agency DFO Fisheries and Oceans Canada EDDMapS Early Detection and Distribution Mapping System F freshwater (salinity) Great Lakes Aquatic Nonindigenous Species Information GLANSIS System GLR Great Lakes region IL Illinois IN Indiana ITIS Integrated Taxonomic Information System M marine (salinity) MB Manitoba MI Michigan MISP Michigan Invasive Species Program MN Minnesota n/a Not available or not applicable NAS Non-indigenous Aquatic Species NY New York NYS New York State OH Ohio OMNRF Ontario Ministry of Natural Resources and Forestry ON Ontario OPIC Ontario Invasive Plant Council PA Pennsylvania PQ Quebec SL standard length TL total length TSN Taxonomic Serial Number US United States USDA United States Department of Agriculture USFWS United States Fish and Wildlife Service USGS United States Geological Survey VT Vermont WI Wisconsin WIDNR Wisconsin Department of Natural Resources

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Appendix B. Current and potential invasive fishes

Table B1. Current and potential invasive fish species in the Great Lakes region, listed alphabetically by scientific name. Taxonomic family and ITIS taxonomic serial number (TSN) are listed for clarity. n/a=data not available.

Scientific name Common name(s) Family ITIS TSN Basis for listing Acanthogobius flavimanus Yellowfin goby Gobiidae 171882 USFWS Alburnus alburnus Common bleak Cyprinidae 163663 Snyder et al. 2014 Alosa aestivalis Blueback herring Clupeidae 161703 Watch list in GLR Alosa maeotica Black sea shad Clupeidae 551289 Snyder et al. 2014 Alosa pseudoharengus Alewife Clupeidae 161706 Regulated in GLR Apeltes quadracus Fourspine stickleback Gasterosteidae 166397 Present in GLR (GLANSIS current) Atherina boyeri Big-scale sand smelt Atherinidae 166024 GLANSIS watch list; Snyder et al. 2014 Babka gymnotrachelus Racer goby Gobiidae 172070 GLANSIS watch list Benthophilus stellatus Starry goby Gobiidae 637036 GLANSIS watch list; Snyder et al. 2014 Carassius auratus Goldfish Cyprinidae 163350 Regulated in GLR Carassius carassius Crusian carp Cyprinidae 163352 Snyder et al. 2014 Carassius gibelio Prussian carp Cyprinidae 688850 USFWS Regulated in GLR, Watch list in GLR; Channa argus Northern snakehead Channidae 166680 Herborg et al. 2007 Regulated in GLR, Watch list in GLR; Channa bleheri Rainbow snakehead Channidae 642749 Herborg et al. 2007 Regulated in GLR, Watch list in GLR; Channa maculata Blotched snakehead Channidae 166684 Herborg et al. 2007 Regulated in GLR, Watch list in GLR; Channa striata Chevron snakehead Channidae 166667 Herborg et al. 2007 Cichlasoma bimaculatum Black acara Cichlidae 169775 USFWS Clarias batrachus Walking catfish Clariidae 164120 Regulated in GLR Clupeonella caspia Caspian tulka Clupeidae n/a Snyder et al. 2014 Clupeonella cultriventris Black, Caspian Sea sprat Clupeidae 161819 GLANSIS watch list; Snyder et al. 2014

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Scientific name Common name(s) Family ITIS TSN Basis for listing Cottus gobio European bullhead Cottidae 167255 Snyder et al. 2014 Ctenopharyngodon idella Grass carp Cyprinidae 163537 Regulated in GLR, Watch list in GLR Cyprinella lutrensis Red shiner Cyprinidae 163792 USGS expanding range in US; CABI Cyprinella whipplei Steelcolor shiner Cyprinidae 163811 GLANSIS watch list Cyprinus carpio Common carp, koi Cyprinidae 163344 Regulated in GLR Fundulus diaphanus Eastern banded killifish, Fundulidae 913978 Regulated in GLR (diaphanus) banded killifish Gambusia affinis Western mosquitofish Poeciliidae 165878 Present in GLR (GLANSIS current) Gambusia holbrooki Eastern mosquitofish Poeciliidae 165896 Regulated in GLR Gasterosteus aculeatus Three spine stickleback Gasterosteidae 201978 Regulated in GLR Gymnocephalus cernuus Ruffe Percidae 168520 Regulated in GLR, Watch list in GLR Hypomesus nipponensis Wakasagi, Japanese smelt Osmeridae 162033 USFWS Hypophthalmichthys harmandi Largescale silver carp Cyprinidae 689232 Regulated in GLR, Watch list in GLR Hypophthalmichthys molitrix Silver carp Cyprinidae 163691 Regulated in GLR, Watch list in GLR Hypophthalmichthys nobilis Bighead carp Cyprinidae 163692 Regulated in GLR, Watch list in GLR Hyrcanogobius bergi Volga dwarf goby Gobiidae 637521 Snyder et al. 2014 GLANSIS watch list, invasive in Southern Ictalurus furcatus Blue catfish Ictaluridae 163997 US Knipowitschia caucasica Caucasian dwarf goby Gobiidae 637541 GLANSIS watch list; Snyder et al. 2014 Lates calcarifer Baramundi Centropomidae 167669 Ontario watch list Leuciscus idus Ide, orfe Cyprinidae 163576 GLANSIS watch list; Snyder et al. 2014 Leuciscus leuciscus Eurasian dace Cyprinidae 163580 GLANSIS watch list; Snyder et al. 2014 Japanese weatherfish, Misgurnus anguillicaudatus Cobitidae 163978 Regulated in GLR weather loach Monopterus albus Asian swamp eel Synbranchidae 166697 USGS NAS: Present in GLR Morone americana White perch Moronidae 167678 Regulated in GLR Morone chrysops x M. saxatilis Wiper, hybrid striped bass Moronidae 167681 Detected in GLR Morone saxatilis Striped bass Moronidae 167680 Status unknown Mylopharyngodon piceus Black carp Cyprinidae 639618 Regulated in GLR, Watch list in GLR

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Scientific name Common name(s) Family ITIS TSN Basis for listing Myxocyprinus asiaticus Chinese hi-fin banded shark Catostomidae 639710 Regulated in GLR Neogobius fluviatilis Monkey goby Gobiidae 172069 Regulated in GLR Neogobius melanostomus Round goby Gobiidae 172072 Regulated in GLR, Watch list in GLR Argentinian silverside, Odontesthes bonariensis Atherinopsidae 630684 USFWS Pejerrey Oncorhynchus keta Chum salmon Salmonidae 161976 GLANSIS, Kolar and Lodge 2002 Oreochromis aureus Blue tilapia Cichlidae 553308 Regulated in GLR Oreochromis niloticus Nile tilapia Cichlidae 553310 Regulated in GLR Osmerus mordax Rainbow smelt Osmeridae 162041 Regulated in GLR Parachanna africana Niger snakehead Channidae 642610 Regulated in GLR; Herborg et al. 2007 Perca fluviatilis Eurasian perch Percidae 168470 Regulated in GLR, Watch list in GLR Perccottus glenii Amur sleeper Odontobutidae 637837 GLANSIS watch list; Pagnucco et al. 2015 Petromyzon marinus Sea lamprey Petromyzontidae 159722 Present, Regulated in GLR Phoxinus phoxinus Eurasian minnow Cyprinidae 163596 Regulated in GLR Poecilia reticulata Guppy Poeciliidae 165903 USFWS Ponticola gorlap Caspian bighead goby Gobiidae n/a Snyder et al. 2014 Proterorhinus semilunaris Freshwater tubenose goby Gobiidae n/a Regulated in GLR, Watch list in GLR Pseudorasbora parva Stone moroko Cyprinidae 689761 USFWS Pterygoplichthys disjunctivus Vermiculated sailfin catfish Loricariidae 680351 USFWS Pterygoplichthys pardalis Amazon sailfin catfish Loricariidae 680353 USFWS Rhodeus ocellatus Rosy bitterling Cyprinidae 689965 USFWS Rhodeus sericeus Amur bitterling Cyprinidae 163607 Regulated in GLR Rutilus rutilus Common roach Cyprinidae 163761 Snyder et al. 2014 Sander lucioperca Zander, pike-perch Percidae 650172 Regulated in GLR, Watch list in GLR Scardinius erythrophthalmus Rudd Cyprinidae 163613 Regulated in GLR, Watch list in GLR Silurus glanis Sheatfish, Wels catfish Siluridae 164068 Watch list in GLR Syngnathus abaster Black-striped pipefish Syngnathidae 166468 Snyder et al. 2014 Tinca tinca Tench Cyprinidae 163348 Regulated in GLR, Watch list in GLR

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Table B2. Geographic ranges and pathways for current and potential invasive fishes in the Great Lakes region (GLR), listed alphabetically by scientific name. Origin (indigenous geographic range), basis for inclusion in list, GLR restrictions, establishment, current ranges, and introduction and spread pathways are listed for each species. n/a=data not available or applicable, italics indicate potential pathways for species yet to arrive in the GLR. GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range North canal, Acanthogobius Northwest America, commercial recreational flavimanus Asia n/a n/a n/a Pacific Asia, shipping boating, bait (Yellowfin goby) Oceania release canal, Alburnus alburnus commercial Europe n/a n/a n/a n/a Europe recreational (Common bleak) shipping boating NY (Finger Atlantic Lakes, canal, Alosa aestivalis North coast, Nova Mohawk North commercial PQ 1995 recreational (Blueback herring) America Scotia to River), VT America shipping boating Florida (Lake Champlain) Ponto- canal, Alosa maeotica commercial Eurasia Caspian n/a n/a n/a Eurasia recreational (Black sea shad) shipping basin boating Atlantic Great Lakes Alosa coast, canal, North (all), North commercial pseudoharengus Labrador to MN 1931 recreational America throughout America shipping (Alewife) South boating GLR Carolina

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Great Lakes (Superior), ON canal, Apeltes quadracus Europe, Europe, (Thunder Bay, North commercial recreational (Fourspine North n/a 1986 Atlantic coast Black and America shipping boating, bait stickleback) America Nipigon release Rivers), PA Ponto- commercial Atherina boyeri Caspian shipping, canal, (Big-scale sand Eurasia basin, n/a n/a n/a Eurasia unauthorized recreational smelt) Mediterranea intentional boating n coast release canal, Babka Ponto- commercial recreational gymnotrachelus Eurasia Caspian n/a n/a n/a Eurasia shipping, boating, bait (Racer goby) basin aquarium/garden release canal, Benthophilus Ponto- commercial recreational stellatus Eurasia Caspian n/a n/a n/a Eurasia shipping, boating, bait (Starry goby) basin aquarium/garden release North America, Great Lakes commercial Africa, aquarium/garden, Carassius auratus (all), shipping, canal, Asia Eastern Asia MN 1879 Eurasia, bait release, (Goldfish) throughout recreational South escaped culture GLR boating America, Oceania canal, recreational Carassius carassius Eurasia, aquarium/garden, Eurasia n/a n/a n/a Europe boating, (Crucian carp) Siberia live trade commercial shipping

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range canal, Carassius gibelio Central North recreational aquarium/garden, (Prussian carp, Eurasia Europe, n/a n/a n/a America, boating, live trade gibel carp) Siberia Eurasia commercial shipping IL, IN, MI, commercial Channa argus North MN, NY, Date aquarium/garden, shipping, canal, (Northern Asia China NY, PA America, OH, ON, unknown live trade recreational snakehead) Asia PQ boating canal, Channa bleheri IL, IN, MI, recreational aquarium/garden, (Rainbow Asia India NY, OH, n/a n/a n/a boating, live trade snakehead) ON, PQ commercial shipping North canal, Channa maculata IL, IN, MI, America recreational Southern aquarium/garden, (Blotched Asia NY, OH, n/a n/a (Hawaii), boating, China live trade snakehead) ON, PQ Africa, commercial Asia shipping wide canal, Channa striata distribution, IL, IN, MI, recreational Asia, aquarium/garden, (Chevron Asia possible NY, OH, n/a n/a boating, Africa live trade snakehead) species ON, PQ commercial complex shipping canal, Cichlasoma recreational South North bimaculatum n/a n/a n/a n/a aquarium/garden boating, America America (Black acara) commercial shipping

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range canal, recreational Clarias batrachus Southeastern North Asia IL, IN, OH n/a n/a aquarium/garden boating, (Walking catfish) Asia America commercial shipping Ponto- canal, Clupeonella caspia commercial Eurasia Caspian n/a n/a n/a Eurasia recreational (Caspian tyulka) shipping basin boating Clupeonella cultriventris Ponto- canal, commercial (Black and Caspian Eurasia Caspian n/a n/a n/a Europe recreational shipping Sea sprat, Black basin boating Sea kilka) canal, Cottus gobio commercial Eurasia Europe, Asia n/a n/a n/a Europe recreational (European bullhead) shipping boating North Mississippi America, IL, IN, MI, commercial Ctenopharyngodon River basin, Africa, MN, NY, Date aquarium/garden, shipping, idella Eurasia Eastern Asia Ohio River, Eurasia, OH, ON, unknown canal, live trade recreational (Grass carp) USA South PQ boating (widespread) America, Oceania recreational Mississippi canal, Cyprinella lutrensis North North boating, bait River basin, n/a n/a n/a commercial (Red shiner) America America release, Mexico shipping aquarium/garden recreational canal, Cyprinella whipplei North Mississippi North boating, bait n/a n/a n/a commercial (Steelcolor shiner) America River basin America release, shipping aquarium/garden

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range North America, Black, Great Lakes commercial Africa, Cyprinus carpio Caspian and (all), aquarium/garden, shipping, Eurasia MN 1880 Eurasia, (Common carp, koi) Aral Sea throughout live trade recreational South basins GLR boating, canal America, Oceania Eastern North Fundulus diaphanus America, Great Lakes commercial North indigenou North recreational (Eastern banded including IL, OH (all), ON, OH, shipping, bait America s America boating, canal killifish) Great Lakes PA release (all, except Superior) North Mississippi America, aquarium/garden, Gambusia affinis River basin, Africa, North Date USA recreational commercial (Western tributaries to n/a Eurasia, America unknown (widespread) boating, canal, shipping mosquitofish) northern Gulf South bait release of Mexico America, Oceania North America, Southeastern aquarium/garden, Gambusia holbrooki Africa, North United Date USA recreational commercial (Eastern IL Eurasia, America States, unknown (widespread) boating, canal, shipping mosquitofish) South Atlantic coast bait release America, Oceania

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Atlantic and Pacific coasts, canal, Gasterosteus including Great Lakes commercial aculeatus aculeatus North Lake Ontario (all), North shipping, IL, OH 1980 aquarium/garden (Threespine America basin (wide throughout America recreational stickleback) distribution, GLR boating, bait possible release species complex) Great Lakes Gymnocephalus IL, IN, MI, North canal, Northern (Superior), commercial cernuus Eurasia MN, NY, 1986 America, recreational Europe, Asia ON, MI, MN, shipping (Ruffe) OH, PQ Eurasia boating WI Hypomesus commercial Japan, North nipponensis shipping, canal, Asia Korean n/a n/a n/a America, unknown (Wakasagi smelt, recreational Peninsula Asia Japanese smelt) boating Hypophthalmichthys commercial harmandi China, aquarium/garden, shipping, canal, Asia IL, MI, PQ n/a n/a n/a (Largescale silver Vietnam live trade recreational carp) boating North America, IL, IN, MI, commercial Hypophthalmichthys Africa, MN, NY, Date Mississippi aquarium/garden, shipping, canal, molitrix Eurasia Eastern Asia Eurasia, OH, ON, unknown River basin, IL live trade recreational (Silver carp) South PQ boating America, Oceania

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range North America, IL, IN, MI, commercial Hypophthalmichthys Asia, MN, NY, Date Mississippi aquarium/garden, shipping, canal, nobilis Asia China Europe, OH, ON, unknown River basin live trade recreational (Bighead carp) South PQ boating America, Oceania commercial Hyrcanogobius Ponto- shipping, canal, bergi Eurasia Caspian n/a n/a n/a Eurasia aquarium/garden recreational (Volga dwarf goby) basin boating, bait release Mississippi commercial North Ictalurus furcatus North River basin, Date aquarium/garden, shipping, canal, n/a IL, OH, MN America, (Blue catfish) America Southern unknown live trade recreational Asia USA, Mexico boating commercial Knipowitschia Ponto- shipping, canal, caucasica Eurasia Caspian n/a n/a n/a Europe aquarium/garden recreational (Caucasian dwarf basin boating, bait goby) release commercial Indo-West Lates calcarifer Asia, aquarium/garden, shipping, canal, Pacific, n/a n/a n/a Asia (Barramundi) Oceania live trade recreational Australia boating commercial North Leuciscus idus Europe, Date aquarium/garden, shipping, canal, Eurasia n/a NY, PA America, (Ide, orfe) Siberia unknown live trade recreational Europe boating

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range canal, Leuciscus leuciscus commercial recreational Eurasia n/a n/a n/a n/a Europe (Eurasian dace) shipping boating, bait release Misgurnus North commercial anguillicaudatus Great Lakes America, aquarium/garden, shipping, canal, (Japanese Asia n/a IL, MI, WI 2005 (MI), IL, MI, Europe, escaped culture recreational weatherfish, NY Oceania boating weather loach) commercial Monopterus albus Asia, Asia, North aquarium/garden, shipping, canal, n/a n/a n/a (Asian swamp eel) Oceania Australia America live trade recreational boating Atlantic Great Lakes coast, St. canal, Morone americana North IL, IN, MN, (all), North commercial Lawrence to 1946 recreational (White perch) America OH throughout America shipping Lake Ontario boating GLR drainages none Southeastern Morone chrysops x (artificial canal, USA Date USA North commercial M. saxatilis hybrid), n/a recreational (artificial unknown (widespread) America shipping (Wiper) North boating hybrid) America North America, St. Lawrence canal, Morone saxatilis North Date Eurasia, commercial Atlantic coast n/a River, USA recreational (Striped bass) America unknown Africa, shipping (widespread) boating South America

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range IL, IN, MI, North commercial Mylopharyngodon MN, NY, Date America, shipping, piceus Asia n/a IL live trade, canal OH, ON, unknown Europe, recreational (Black carp) PQ Asia boating Myxocyprinus commercial asiaticus shipping, canal, Asia China IL n/a n/a Asia aquarium/garden (Chinese hi-fin recreational banded shark) boating Ponto- canal, commercial Neogobius fluviatilis Caspian recreational Eurasia IL n/a n/a Europe shipping, (Monkey goby) basin, boating, bait aquarium/garden Turkey release commercial Great Lakes Neogobius Ponto- IL, IN, MI, North commercial shipping, (all), melanostomus Eurasia Caspian MN, NY, 1990 America, shipping, recreational throughout (Round goby) basin OH, PQ Europe aquarium/garden boating, bait GLR release Odontesthes Southern Europe, commercial bonariensis South Argentina, South shipping, canal, n/a n/a n/a unknown (Argentinian America Rio de La America, recreational silverside) Plata Asia boating commercial North Oncorhynchus keta North shipping, canal, America, North Pacific n/a n/a n/a unknown (Chum salmon) America recreational Asia boating

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range North America, Africa, commercial Oreochromis aureus Africa, Asia, aquarium/garden, shipping, canal, Africa IL, MN n/a n/a (Blue tilapia) Middle East Europe, live trade recreational Central boating America, Oceania commercial Oreochromis North aquarium/garden, shipping, canal, niloticus Africa n/a IL, MN n/a n/a America, live trade recreational (Nile tilapia) Africa boating Great Lakes commercial North North Pacific, intentional Osmerus mordax (all), North shipping, America, North MN 1923 release, bait (Rainbow smelt) throughout America recreational Asia Atlantic release GLR boating commercial Parachanna IL, IN, MI, Benin, aquarium/garden, shipping, canal, africana Africa NY, OH, n/a n/a n/a Nigeria live trade recreational (Niger snakehead) ON boating North America, canal, Perca fluviatilis Africa, commercial Eurasia n/a IL, PQ n/a n/a recreational (Eurasian perch) Oceania, shipping boating Europe, Asia commercial Perccottus glenii Northeastern shipping, canal, Asia n/a n/a n/a Europe aquarium/garden (Amur sleeper) Asia recreational boating

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range North Great Lakes Petromyzon canal, America, Atlantic (all), North commercial marinus IL, MN, OH 1921 recreational Europe, Coast throughout America shipping (Sea lamprey) boating Africa GLR canal, Phoxinus phoxinus commercial recreational Eurasia n/a IL n/a n/a Eurasia (Eurasian minnow) shipping boating, bait release canal, Asia, commercial Africa, Poecilia reticulata South shipping, n/a n/a n/a n/a Oceania, aquarium/garden (Guppy) America recreational North boating, bait America release canal, Ponticola gorlap Ponto- commercial recreational (Caspian bighead Eurasia Caspian n/a n/a n/a Eurasia shipping, boating, bait goby) basin aquarium/garden release Great Lakes Proterorhinus IL, IN, MI, (Erie), St. commercial recreational semilunaris Black Sea North Eurasia MN, OH, 1990 Claire River, shipping, boating, bait (Freshwater basin America PQ Lake St. aquarium/garden release tubenose goby) Claire commercial Pseudorasbora Eurasia, shipping, canal, parva Asia East Asia ON n/a n/a aquarium/garden Oceania recreational (Stone moroko) boating Pterygoplichthys commercial disjunctivus South Amazon North shipping, canal, n/a n/a n/a aquarium/garden (Vermiculated sailfin America River basin America recreational catfish) boating

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Pterygoplichthys commercial North pardalis South Amazon shipping, canal, n/a n/a n/a America, aquarium/garden (Amazon sailfin America River basin recreational Asia catfish) boating commercial Rhodeus ocellatus Eastern Asia, shipping, canal, Asia n/a n/a n/a Eurasia aquarium/garden (Rosy bitterling) Taiwan recreational boating commercial Amur River North Rhodeus sericeus Date NY (Bronx shipping, Eurasia basin, IL, MI America, aquarium/garden (Amur bitterling) unknown River) recreational Europe Europe boating Ponto- commercial Rutilus rutilus Caspian shipping, canal, Eurasia n/a n/a n/a Europe unknown (Common roach) basin, recreational Europe boating North commercial Sander lucioperca Europe, IL, IN, MN, America, shipping, canal, Eurasia n/a n/a unknown (Zander, pikeperch) Siberia PQ, ON Asia, recreational Europe boating intentional Great Lakes North release, bait Scardinius Europe, IL, IN, MI, (Erie, America, release, commercial erythrophthalmus Eurasia Western MN, NY, 1990 Ontario), St. Europe, recreational shipping, canal (Rudd) Asia, Siberia OH, PQ Lawrence Africa, boating, escaped River, ON, NY Oceania culture commercial Silurus glanis Europe, Europe, shipping, canal, (Sheatfish, Wels Eurasia ON, PQ n/a n/a aquarium/garden Western Asia Africa recreational catfish) boating

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Syngnathus abaster (Black-striped commercial canal, Eastern pipefish, Eurasia n/a n/a n/a Eurasia shipping, recreational Atlantic shortsnouted aquarium/garden boating pipefish) North Mississippi America, commercial River basin, Africa, intentional Tinca tinca Europe, Date shipping, canal, Eurasia IN, MI, PQ PQ (Richelieu Eurasia, release, escaped (Tench) Western Asia unknown recreational River), USA South culture boating (widespread) America, Oceania

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Table B3. Size and habitat preferences for current and potential invasive fishes to the Great Lakes region, listed alphabetically by scientific name. n/a=data not available; TL=total length; SL=standard length; FL=fork length. Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) Acanthogobius freshwater, lake/pond, flavimanus 30 n/a 3 cold brackish, benthic, shallow, soft substrates river/stream (Yellowfin goby) marine Alburnus alburnus freshwater, lake/pond, 25 60 n/a cool benthopelagic (Common bleak) brackish river/stream freshwater, Alosa aestivalis lake/pond, 40 n/a 8 cold brackish, pelagic (Blueback herring) river/stream marine freshwater, Alosa maeotica lake/pond, 32 n/a 6 cool brackish, pelagic (Black sea shad) river/stream marine Alosa freshwater, lake/pond, pseudoharengus 25 200 5–11 cool brackish, pelagic river/stream (Alewife) marine Apeltes quadracus freshwater, lake/pond, (Fourspine 6.4 n/a 2–3 brackish, benthopelagic, littoral river/stream stickleback) marine Atherina boyeri freshwater, (Big-scale sand 15 n/a 4 cold brackish, lake/pond pelagic, benthic smelt) marine Babka benthopelagic, fast to still water with freshwater, lake/pond, gymnotrachelus 16 TL n/a 5 cold sand, mud, gravel substrates, littoral brackish river/stream (Racer goby) vegetation or cryptic habitats Benthophilus freshwater, benthic, muddy substrates more than stellatus 13.5 TL n/a 3 cold river/stream brackish 3 m deep (Starry goby)

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) lake/pond, Carassius auratus freshwater, benthopelagic, still to slow-moving 40 TL 300 6–41 warm river/stream, (Goldfish) brackish eutrophic water, littoral vegetation wetland benthic, still to slow-moving water, Carassius carassius freshwater, lake/pond, 50 5000 10 warm prefers dense vegetation, burrows in (Crucian carp) brackish river/stream mud (dry season, winter) Carassius gibelio benthopelagic, still to slow-moving freshwater, lake/pond, (Prussian carp, gibel 46 3000 10 cold water, prefers dense vegetation, brackish river/stream carp) eutrophic water benthopelagic, prefers still to slow- Channa argus lake/pond, moving, shallow water with mud (Northern 85 8000 8 warm freshwater river/stream, substrate or vegetation, obligate air snakehead) wetland breather survives out of water for up to 4 days benthopelagic, prefers still to slow- Channa bleheri lake/pond, moving, shallow water with mud (Rainbow 20 n/a n/a cool freshwater river/stream, substrate or vegetation, facultative air snakehead) wetland breather Channa maculata lake/pond, benthopelagic, prefers shallow water (Blotched 33 TL n/a n/a warm freshwater river/stream, with mud substrate or vegetation, snakehead) wetland facultative air breather benthopelagic, prefers stagnant, lake/pond, Channa striata freshwater, shallow and muddy water, survives 100 TL 3000 n/a cool river/stream, (Chevron snakehead) brackish dry season by burrowing in mud, wetland obligate air breather Cichlasoma river/stream, bimaculatum 12 SL n/a n/a warm freshwater benthopelagic wetland (Black acara)

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) benthic, prefers stagnant, muddy or lake/pond, Clarias batrachus freshwater, swampy water with high turbidity, 47 TL 1200 5–25 warm river/stream, (Walking catfish) brackish survives dry season by burrowing in wetland mud, facultative air breather freshwater, Clupeonella caspia lake/pond, 12 SL n/a n/a cold brackish, pelagic (Caspian tyulka) river/stream marine Clupeonella cultriventris freshwater, lake/pond, (Black and Caspian 12.5 SL n/a 1–6 cool brackish, pelagic river/stream Sea sprat, Black Sea marine kilka) benthic, prefers cold, clear and fast Cottus gobio freshwater, lake/pond, 18 TL n/a 4–10 cold flowing water, sand, gravel or rocky (European bullhead) brackish river/stream shores Ctenopharyngodon freshwater, lake/pond, idella 125 n/a 5–33 warm benthic, still water with vegetation brackish river/stream (Grass carp) benthopelagic, silty, sandy and rocky Cyprinella lutrensis 9 n/a 3 warm freshwater river/stream pools and runs (sometimes riffles) of (Red shiner) creeks to medium rivers benthopelagic, rocky and sandy runs, Cyprinella whipplei 16 TL n/a 3–4 cool freshwater river/stream riffles and pools of creeks to medium (Steelcolor shiner) rivers with permanent flow benthopelagic, slow and still water, Cyprinus carpio freshwater, lake/pond, 120 TL n/a 13–38 warm prefers submerged vegetation and soft (Common carp, koi) brackish river/stream substrates, thrives in turbidity benthopelagic, shallow, slow and still Fundulus diaphanus freshwater, lake/pond, water with soft substrates (sand, (Eastern banded 13 n/a 3–4 cool brackish river/stream gravel or detritus) and submerged killifish) vegetation

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) Gambusia affinis freshwater, lake/pond, benthopelagic, prefers standing to (Western 7 TL n/a 3 warm brackish, river/stream, slow water with submerged vegetation mosquitofish) marine wetland and soft substrates Gambusia holbrooki freshwater, lake/pond, benthopelagic, prefers standing to (Eastern 8 TL n/a 1–2 warm brackish, river/stream, slow water with submerged vegetation mosquitofish) marine wetland and soft substrates benthopelagic, two freshwater Gasterosteus morphological variations: limnetic freshwater, lake/pond, aculeatus aculeatus (prefers oligotrophic lakes) and 8 TL n/a 1–3 cold brackish, river/stream, (Threespine benthic (prefers shallow eutrophic marine wetland stickleback) water with soft substrates and vegetation) benthic, eutrophic slow or still water Gymnocephalus freshwater, lake/pond, with soft substrates and absent of cernuus 25 400 10 cool brackish river/stream vegetation or deep water with sand or (Ruffe) gravel Hypomesus freshwater, nipponensis 17 TL n/a n/a brackish, lake/pond pelagic (Wakasagi smelt, marine Japanese smelt) Hypophthalmichthys benthopelagic, prefers slow or still harmandi freshwater, lake/pond, 55 SL n/a 20 cool shallow water, large rivers with (Largescale silver brackish river/stream marked water level fluctuations carp) Hypophthalmichthys benthopelagic, prefers slow or still freshwater, lake/pond, molitrix 105 TL 50,000 10–20 warm shallow water, large rivers with brackish river/stream (Silver carp) marked water level fluctuations Hypophthalmichthys benthopelagic, prefers slow or still lake/pond, nobilis 146 SL 40,000 10–33 warm freshwater shallow water, large rivers with river/stream (Bighead carp) marked water level fluctuations

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) freshwater, Hyrcanogobius bergi lake/pond, benthopelagic, prefers soft substrates 3.6 SL n/a 1 cold brackish, (Volga dwarf goby) river/stream (mud, sand) marine benthopelagic, variety of habitats, Ictalurus furcatus freshwater, lake/pond, prefers clear, strong flowing water and 165 TL 68,000 9–25 cool (Blue catfish) brackish river/stream harder substrates, as well as still water with softer substrates Knipowitschia freshwater, lake/pond, caucasica benthic, prefers shallow, soft 5 TL n/a 1–2 cold brackish, river/stream, (Caucasian dwarf substrates and vegetation marine wetland goby) freshwater, pelagic, clear to turbid water, prefer Lates calcarifer lake/pond, 200 TL 60,000 20 warm brackish, cover (banks, logs, overhanging (Barramundi) river/stream marine vegetation) Leuciscus idus freshwater, lake/pond, benthopelagic, prefers slow or still 102 4000 18 cold (Ide, orfe) brackish river/stream eutrophic water benthopelagic, prefers moderate to Leuciscus leuciscus freshwater, lake/pond, 40 TL 1000 16 cold/cool fast flowing water with hard substrates (Eurasian dace) brackish river/stream (rock, gravel) Misgurnus anguillicaudatus lake/pond, benthic, prefers muddy substrates, (Japanese 28 SL n/a 10 cold freshwater river/stream, leaf litter and aquatic macrophyte weatherfish, weather wetland beds loach) lake/pond, benthic, prefers slow-moving water, Monopterus albus freshwater, 100 SL n/a n/a warm river/stream, survives dry season by burrowing into (Asian swamp eel) brackish wetland moist earth freshwater, lake/pond, Morone americana benthic, prefers quiet water and soft 58 TL 2200 16 warm brackish, river/stream, (White perch) substrate (mud) marine wetland

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) Morone chrysops x lake/pond, benthic; pelagic, prefers slow to still M. saxatilis 90 11,000 5-8 cool freshwater river/stream water (Wiper) freshwater, Morone saxatilis lake/pond, benthic; pelagic, prefers slow to still 200 TL 57,000 10-30 cool brackish, (Striped bass) river/stream water marine Mylopharyngodon lake/pond, piceus 180 TL 70,000 15 warm freshwater benthic, prefers clear water river/stream (Black carp) Myxocyprinus asiaticus lake/pond, 60 TL n/a n/a cold freshwater benthic (Chinese hi-fin river/stream banded shark) Neogobius fluviatilis freshwater, lake/pond, benthopelagic, still to slow water with 20 SL n/a 5 warm (Monkey goby) brackish river/stream soft substrates (sand, mud) Neogobius freshwater, benthic, prefers shallow water with lake/pond, melanostomus 25 TL n/a 6 warm brackish, hard or soft substrates, submerged river/stream (Round goby) marine vegetation Odontesthes freshwater, bonariensis lake/pond, 50 TL n/a 7 cold/cool brackish, pelagic (Argentinian river/stream marine silverside) freshwater, Oncorhynchus keta lake/pond, 100 FL 15,900 7 cold brackish, benthopelagic (Chum salmon) river/stream marine Oreochromis aureus freshwater, lake/pond, benthopelagic, prefers open water or 51 2000 5 warm (Blue tilapia) brackish river/stream among stones and vegetation Oreochromis niloticus freshwater, lake/pond, 60 SL 4300 9 warm benthopelagic, prefers shallow water (Nile tilapia) brackish river/stream

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) freshwater, Osmerus mordax lake/pond, pelagic, prefers still water of lakes and 36 TL n/a 7 cold brackish, (Rainbow smelt) river/stream medium to large rivers marine Parachanna africana lake/pond, 17 SL n/a n/a warm freshwater benthopelagic, facultative air breather (Niger snakehead) river/stream benthopelagic, lakes of all types to Perca fluviatilis freshwater, lake/pond, 60 SL 4800 22 cool medium sized streams, prefers clear (Eurasian perch) brackish river/stream water benthic, prefers slow to still shallow lake/pond, water with soft substrates and Perccottus glenii freshwater, 25 TL 250 4–10 cool river/stream, vegetation, survives dry season by (Amur sleeper) brackish wetland burrowing in mud, facultative air breather freshwater, Petromyzon marinus lake/pond, 120 TL 2500 5–9 cold brackish, benthic (Sea lamprey) river/stream marine benthic, wide habitat range, able to thrive in fast, slow and still oligotrophic Phoxinus phoxinus freshwater, lake/pond, 14 TL n/a 6–11 cold water, prefers shallow lakes and slow (Eurasian minnow) brackish river/stream flowing streams, hard substrates and vegetation benthopelagic, wide habitat range Poecilia reticulata freshwater, lake/pond, 5 SL n/a 3–5 cool/warm from highly turbid still water to clear (Guppy) brackish river/stream streams Ponticola gorlap freshwater, benthic, prefers slow or still water with lake/pond, (Caspian bighead 33 TL n/a n/a cold brackish, sand or rock substrate, littoral river/stream goby) marine vegetation Proterorhinus benthopelagic, prefers slow or still semilunaris freshwater, lake/pond, 12 TL n/a 5 cold water with dense vegetation, (Freshwater brackish river/stream substrates of coarse rock or sand tubenose goby)

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) Pseudorasbora parva lake/pond, benthopelagic, wide habitat range, 11 TL n/a 3–5 cold freshwater (Stone moroko) river/stream prefers submerged vegetation Pterygoplichthys lake/pond, disjunctivus freshwater, benthic, wide habitat range, facultative 70 TL n/a 5–10 cool river/stream, (Vermiculated sailfin brackish air breather wetland catfish) Pterygoplichthys lake/pond, pardalis freshwater, benthic, wide habitat range, facultative 50 TL 310 5 warm river/stream, (Amazon sailfin brackish air breather wetland catfish) benthopelagic, prefers still to slow- lake/pond, Rhodeus ocellatus moving shallow water with dense 5.8 SL n/a 3–4 warm freshwater river/stream, (Rosy bitterling) vegetation and soft substrates (sand, wetland silt) lake/pond, Rhodeus sericeus 11 TL n/a 5 cool freshwater river/stream, benthopelagic, prefers soft substrates (Amur bitterling) wetland benthopelagic, wide habitat range, Rutilus rutilus freshwater, lake/pond, 50 TL 1800 14 cool most abundant in eutrophic lakes and (Common roach) brackish river/stream medium to large rivers Sander lucioperca freshwater, lake/pond, pelagic, prefers slow or still eutrophic, 100 SL 20,000 8–24 cool (Zander, pikeperch) brackish river/stream turbid, well oxygenated water Scardinius benthopelagic, wide habitat range, freshwater, lake/pond, erythrophthalmus 48 TL n/a 15–19 cold prefers slow or still eutrophic, well brackish river/stream (Rudd) vegetated water benthopelagic; benthic, wide habitat Silurus glanis freshwater, lake/pond, range, prefers large slow or still water (Sheatfish, Wels 500 TL 306,000 15–30 cool brackish river/stream with vegetation and cryptic habitat, catfish) tolerates eutrophic and turbid water

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Maximum Maximum Species adult adult Longevity Thermal Salinity Habitat Habitat description (common name) length weight (years) guild tolerance preference (cm) (g) Syngnathus abaster (Black-striped freshwater, benthic, prefers slow to still water with lake/pond, pipefish, 21 TL n/a 4 cold brackish, soft substrate (sand, mud) with cryptic river/stream shortsnouted marine habitats (detritus, vegetation) pipefish) Tinca tinca freshwater, lake/pond, benthic, prefers slow or still water with 84 TL 7500 20 cool (Tench) brackish river/stream soft substrates (mud) and vegetation

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Table B4. Diet and reproductive behaviour for current and potential invasive fishes to the Great Lakes region, listed alphabetically by scientific name. n/a=data not available. Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Acanthogobius sexual; guarder, nest flavimanus carnivore benthic invertebrates, small fish 1 amphidromous spawner (Yellowfin goby) Alburnus alburnus sexual; nonguarder, carnivore zooplankton, benthic macroinvertebrates 2–3 (Common bleak) potamodromous broadcast spawner Alosa aestivalis sexual; nonguarder, carnivore zooplankton, small fish 3–4 (Blueback herring) anadromous broadcast spawner Alosa maeotica sexual; nonguarder, carnivore zooplankton, small fish 2 (Black sea shad) anadromous broadcast spawner 1–3 Alosa (landlocked); sexual; nonguarder, pseudoharengus omnivore phytoplankton, zooplankton, small fish 3–5 anadromous broadcast spawner (Alewife) (anadromous) Apeltes quadracus sexual; guarder, nest (Fourspine carnivore zooplankton, benthic macroinvertebrates 1 migratory spawner stickleback) Atherina boyeri zooplankton, benthic macroinvertebrates, sexual; nonguarder, (Big-scale sand carnivore 1 small fish amphidromous broadcast spawner smelt) Babka sexual; non- guarder, nest gymnotrachelus carnivore benthic macroinvertebrates, small fish 2 migratory spawner (Racer goby) Benthophilus sexual; non- guarder, substrate stellatus carnivore benthic macroinvertebrates, small fish 3 migratory chooser (Starry goby) Carassius auratus benthic macroinvertebrates, plant material, sexual; nonguarder, omnivore 1 (Goldfish) algae, zooplankton, detritus, fish eggs/fry potamodromous broadcast spawner Carassius carassius benthic macroinvertebrates, zooplankton, sexual; nonguarder, omnivore 1–3 (Crucian carp) algae, plant material, detritus potamodromous broadcast spawner

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Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Carassius gibelio benthic macroinvertebrates, zooplankton, sexual, asexual; nonguarder, (Prussian carp, gibel omnivore 2 algae, plant material, detritus potamodromous broadcast spawner carp) Channa argus zooplankton, benthic macroinvertebrates, sexual; non- guarder, substrate (Northern carnivore small fish, fish, insects, amphibians, small 2–3 migratory chooser snakehead) mammals Channa bleheri zooplankton, benthic macroinvertebrates, sexual; non- guarder, substrate (Rainbow carnivore n/a small fish, fish, insects, amphibians migratory chooser snakehead) Channa maculata zooplankton, benthic macroinvertebrates, sexual; non- guarder, substrate (Blotched carnivore n/a small fish, fish, insects, amphibians migratory chooser snakehead) Channa striata zooplankton, benthic macroinvertebrates, sexual; guarder, substrate carnivore n/a (Chevron snakehead) small fish, fish, insects, amphibians potamodromous chooser Cichlasoma benthic macroinvertebrates, insects, small sexual; non- guarder, substrate bimaculatum omnivore n/a fish, fish eggs migratory chooser (Black acara) Clarias batrachus detritus, benthic macroinvertebrates, sexual; guarder, nest omnivore 1 (Walking catfish) amphibians, fish potamodromous spawner Clupeonella caspia sexual; nonguarder, carnivore zooplankton 1 (Caspian tyulka) anadromous broadcast spawner Clupeonella cultriventris sexual; nonguarder, (Black and Caspian carnivore zooplankton, small fish 1 anadromous broadcast spawner Sea sprat, Black Sea kilka) Cottus gobio sexual; guarder, nest carnivore benthic invertebrates, small fish 1–3 (European bullhead) potamodromous spawner Ctenopharyngodon submerged grasses, aquatic macrophytes, sexual; nonguarder, idella omnivore 2–10 detritus, insects, benthic macroinvertebrates potamodromous broadcast spawner (Grass carp)

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Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Cyprinella lutrensis insects, benthic macroinvertebrates, algae, sexual; non- nonguarder, brood omnivore 1–2 (Red shiner) macrophytes, fish eggs, small fish migratory hider Cyprinella whipplei insects, zooplankton, benthic invertebrates, sexual; non- nonguarder, brood omnivore 1–2 (Steelcolor shiner) detritus migratory hider Cyprinus carpio plant material, detritus, zooplankton, benthic sexual; nonguarder, omnivore 3–5 (Common carp, koi) invertebrates, fish eggs potamodromous broadcast spawner Fundulus diaphanus generalist omnivore, benthic invertebrates, sexual; non- nonguarder, (Eastern banded omnivore 1 insects, zooplankton migratory broadcast spawner killifish) Gambusia affinis zooplankton, benthic macroinvertebrates, sexual; (Western carnivore insects, small fish, fish, fish eggs, 1–3 bearer, internal potamodromous mosquitofish) amphibians, detritus Gambusia holbrooki zooplankton, benthic macroinvertebrates, sexual; (Eastern carnivore insects, small fish, fish, fish eggs, 1–3 bearer, internal potamodromous mosquitofish) amphibians, detritus Gasterosteus aculeatus aculeatus zooplankton, benthic macroinvertebrates, sexual; guarder, nest carnivore 1–2 (Threespine fish eggs, small fish anadromous spawner stickleback) Gymnocephalus zooplankton, benthic macroinvertebrates, sexual; nonguarder, cernuus carnivore 1 fish eggs, small fish potamodromous broadcast spawner (Ruffe) Hypomesus nipponensis zooplankton, benthic macroinvertebrates, sexual; nonguarder, carnivore 1–2 (Wakasagi smelt, insects anadromous broadcast spawner Japanese smelt) Hypophthalmichthys harmandi sexual; nonguarder, omnivore zooplankton, phytoplankton 5–6 (Largescale silver potamodromous broadcast spawner carp)

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Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Hypophthalmichthys zooplankton, phytoplankton, detritus, sexual; nonguarder, molitrix omnivore 4–6 bacteria potamodromous broadcast spawner (Silver carp) Hypophthalmichthys zooplankton, phytoplankton, benthic algae, sexual; nonguarder, nobilis omnivore 2–3 detritus potamodromous broadcast spawner (Bighead carp) Hyrcanogobius bergi guarder, nest carnivore benthic macroinvertebrates, zooplankton 1 sexual; n/a (Volga dwarf goby) spawner Ictalurus furcatus fish, benthic macroinvertebrates, detritus, sexual; non- guarder, nest omnivore 4–7 (Blue catfish) amphibians, small mammals, birds migratory spawner Knipowitschia caucasica sexual; guarder, nest carnivore benthic macroinvertebrates, zooplankton 1 (Caucasian dwarf amphidromous spawner goby) sexual Lates calcarifer fish, small fish, fish eggs, benthic nonguarder, carnivore 2–8 (protandrous); (Barramundi) macroinvertebrates broadcast spawner catadromous Leuciscus idus fish, small fish, benthic invertebrates, sexual; nonguarder, omnivore 3–5 (Ide, orfe) macrophytes, insects potamodromous broadcast spawner Leuciscus leuciscus macrophytes, algae, zooplankton, benthic sexual; nonguarder, omnivore 3-4 (Eurasian dace) macroinvertebrates, detritus potamodromous broadcast spawner Misgurnus anguillicaudatus benthic macroinvertebrates, detritus, sexual; non- nonguarder, (Japanese carnivore 1–2 zooplankton migratory broadcast spawner weatherfish, weather loach) sexual Monopterus albus fish, fish eggs, benthic macroinvertebrates, guarder, nest carnivore n/a (protandrous); (Asian swamp eel) detritus, amphibians spawner potamodromous Morone americana benthic macroinvertebrates, fish eggs, small sexual; nonguarder, carnivore 2–3 (White perch) fish anadromous broadcast spawner

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Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Morone chrysops x opportunistic omnivore, fish, benthic sexual; nonguarder, M. saxatilis omnivore 2–3 macroinvertebrates, fish eggs, small fish potamodromous broadcast spawner (Wiper) Morone saxatilis opportunistic omnivore, fish, benthic sexual; nonguarder, omnivore 2–6 (Striped bass) macroinvertebrates, fish eggs, small fish anadromous broadcast spawner Mylopharyngodon benthic macroinvertebrates (adults: primarily sexual; nonguarder, piceus carnivore 4–6 molluscs), zooplankton potamodromous broadcast spawner (Black carp) Myxocyprinus asiaticus benthic macroinvertebrates, algae, detritus, sexual; nonguarder, omnivore n/a (Chinese hi-fin plant material potamodromous broadcast spawner banded shark) Neogobius fluviatilis benthic macroinvertebrates, amphibians, guarder, substrate carnivore 2 sexual; n/a (Monkey goby) fish, zooplankton chooser Neogobius benthic macroinvertebrates (including sexual; guarder, substrate melanostomus omnivore 2–4 molluscs), zooplankton, small fish, fish eggs amphidromous chooser (Round goby) Odontesthes bonariensis zooplankton, phytoplankton, benthic algae, sexual; non- nonguarder, omnivore 1 (Argentinian fish, benthic macroinvertebrates migratory broadcast spawner silverside) Oncorhynchus keta opportunistic omnivore, zooplankton, fish, sexual; nonguarder, brood omnivore 2–4 (Chum salmon) benthic macroinvertebrates anadromous hider phytoplankton, benthic algae, zooplankton, Oreochromis aureus omnivore, sexual; benthic invertebrates, small fish, detritus, 0.5 bearer, external (Blue tilapia) herbivore potamodromous plant material phytoplankton, benthic algae, detritus, Oreochromis niloticus omnivore, sexual; zooplankton, benthic macroinvertebrates, 0.5 bearer, external (Nile tilapia) herbivore potamodromous fish eggs Osmerus mordax zooplankton, small fish, fish eggs, benthic sexual; nonguarder, carnivore 2–3 (Rainbow smelt) macroinvertebrates anadromous broadcast spawner

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Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Parachanna africana sexual; guarder, substrate carnivore fish, benthic macroinvertebrates, amphibians n/a (Niger snakehead) potamodromous chooser Perca fluviatilis opportunistic omnivore, zooplankton, fish, sexual; nonguarder, omnivore 1–4 (Eurasian perch) benthic macroinvertebrates anadromous broadcast spawner opportunistic omnivore, benthic Perccottus glenii guarder, nest omnivore macroinvertebrates, amphibians, small fish, 1–3 sexual; n/a (Amur sleeper) spawner fish eggs, macrophytes adults are parasitic (fish, outside host), Petromyzon marinus endoparasite sexual; nonguarder, brood juveniles prefer zooplankton, detritus, fish 3–5 (Sea lamprey) , carnivore anadromous hider (parasitic) Phoxinus phoxinus algae, detritus, benthic macroinvertebrates, sexual; nonguarder, omnivore 2–4 (Eurasian minnow) insects, plant material, zooplankton potamodromous broadcast spawner Poecilia reticulata algae, zooplankton, benthic sexual; non- omnivore 0.08–0.25 bearer, internal (Guppy) macroinvertebrates, detritus migratory Ponticola gorlap guarder, substrate (Caspian bighead carnivore benthic macroinvertebrates, small fish 1–2 sexual; n/a chooser goby) Proterorhinus semilunaris benthic macroinvertebrates, zooplankton, guarder, nest carnivore 1–2 sexual; n/a (Freshwater small fish spawner tubenose goby) zooplankton, benthic macroinvertebrates, Pseudorasbora parva sexual; non- nonguarder, brood omnivore small fish, fish eggs, plant material, 1 (Stone moroko) migratory hider phytoplankton Pterygoplichthys disjunctivus detritus, algae, benthic macroinvertebrates, guarder, nest omnivore n/a sexual; n/a (Vermiculated sailfin fish eggs spawner catfish) Pterygoplichthys pardalis detritus, algae, benthic macroinvertebrates, guarder, nest omnivore n/a sexual; n/a (Amazon sailfin fish eggs spawner catfish)

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Age to Species Trophic Diet details maturity Reproduction Reproduction guild (common name) guild (years) Rhodeus ocellatus sexual; nonguarder, brood omnivore zooplankton, phytoplankton 1 (Rosy bitterling) potamodromous hider Rhodeus sericeus algae, phytoplankton, detritus, benthic sexual; nonguarder, brood omnivore 1 (Amur bitterling) macroinvertebrates potamodromous hider opportunistic omnivore, shifts between littoral and pelagic, benthic Rutilus rutilus sexual; nonguarder, omnivore macroinvertebrates (including molluscs), 2–4 (Common roach) potamodromous broadcast spawner zooplankton, phytoplankton, benthic algae, benthic macrophytes, plant material, detritus Sander lucioperca primarily piscivorous, fish, zooplankton, sexual; guarder, nest carnivore 2–3 (Zander, Pikeperch) benthic macroinvertebrates potamodromous spawner Scardinius macrophytes, algae, zooplankton, benthic omnivore, sexual; nonguarder, erythrophthalmus macroinvertebrates, detritus, small fish, fish 2–3 herbivore potamodromous broadcast spawner (Rudd) eggs Silurus glanis opportunistic forager, benthic sexual; non- guarder, nest (Sheatfish, Wels omnivore macroinvertebrates, fish, birds, small 3–4 migratory spawner catfish) mammals Syngnathus abaster (Black-striped sexual; pipefish, carnivore small fish, macroinvertebrates 1 bearer, external amphidromous Shortsnouted pipefish) Tinca tinca benthic macroinvertebrates (including sexual; nonguarder, omnivore 2–7 (Tench) molluscs), detritus, plant material potamodromous broadcast spawner

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Table B5. Impacts of invasion (realized or potential) for current and potential invasive fishes in the Great Lakes region, listed alphabetically by scientific name. n/a=data not availalble. Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment Acanthogobius alters indigenous fish communities, competition, flavimanus n/a n/a n/a competes with indigenous communities USFWS predation (Yellowfin goby) for food and resources competes with indigenous communities Alburnus alburnus little/no competition, high high for food and resources, may cause USFWS (Common bleak) evidence ecosystem change ecosystem changes Alosa aestivalis inadequate little/no little/no alters food web; predation on competition n/a (Blueback herring) research evidence evidence zooplankton communities Alosa maeotica competes with indigenous communities n/a n/a n/a competition n/a (Black sea shad) for food and resources Alosa competition, alters lake food web by heavy pseudoharengus high high high USFWS predation zooplankton predation (Alewife) Apeltes quadracus competes with indigenous communities inadequate little/no inadequate (Fourspine competition for food and resources, displaces USFWS research evidence research stickleback) indigenous sticklebacks competes with indigenous communities Atherina boyeri little/no for food and resources, able to reach (Big-scale sand moderate high competition USFWS evidence high densities quickly, predation on smelt) zooplankton communities Babka competes with indigenous communities inadequate little/no inadequate competition, gymnotrachelus for food and resources; fish parasite USFWS research evidence research disease vector (Racer goby) host Benthophilus inadequate little/no little/no may compete with indigenous species stellatus competition USFWS research evidence evidence for food and resources (Starry goby)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment alters indigenous fish communities and competition, food webs, competes with indigenous Carassius auratus inadequate little/no inadequate predation, communities for food and resources, NYS (Goldfish) research evidence research ecosystem change increases risk of algal blooms, increases water turbidity weak competitor, absent in waters with Carassius carassius inadequate little/no rich fish communities and predatory moderate not likely USFWS (Crucian carp) research evidence species, abundant in the absence of other fish species dominant species, competes with indigenous communities for food and Carassius gibelio resources, alters indigenous inadequate little/no inadequate competition, (Prussian carp, gibel communities and food webs, may cause USFWS research evidence research ecosystem change carp) ecosystem changes, increases water turbidity, competes primarily with indigenous cyprinids dominant predator competes with Channa argus competition, indigenous communities for food and inadequate DFO, (Northern moderate moderate predation, disease resources, heavy predation on research USFWS snakehead) vector indigenous prey communities, parasite and pathogen host Channa bleheri competes with indigenous communities competition, (Rainbow n/a n/a n/a for food and resources, likely predation n/a predation snakehead) on indigenous communities dominant predator competes with Channa maculata competition, indigenous communities for food and (Blotched n/a n/a n/a n/a predation resources, likely predation on snakehead) indigenous communities dominant predator competes with Channa striata competition, indigenous communities for food and (Chevron n/a n/a n/a n/a predation resources, likely predation on snakehead) indigenous communities

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous communities Cichlasoma competition, (sunfish) for food and resources bimaculatum n/a n/a n/a USFWS predation (spawning sites), predation on (Black acara) indigenous communities dominant predator competes with indigenous communities (especially Clarias batrachus competition, n/a n/a n/a centrarchids and catfish) for food and n/a (Walking catfish) predation resources, likely predation on indigenous communities Clupeonella caspia competes with indigenous fish for food n/a n/a n/a competition n/a (Caspian tyulka) and resources Clupeonella cultriventris inadequate little/no little/no competes with indigenous fish for food (Black and Caspian competition USFWS research evidence evidence and resources Sea sprat, Black Sea kilka) Cottus gobio inadequate little/no competes with indigenous fish for food n/a competition USFWS (European bullhead) research evidence and resources competes with indigenous fish for food Ctenopharyngodon competition, and resources, alteration of indigenous DFO, idella high low high ecosystem change, habitat and food webs (consumes large USFWS (Grass carp) disease vector amounts of macrophytes), parasitic tapeworm and viral host competes with indigenous fish competition, Cyprinella lutrensis (especially Cyprinids) for food and n/a n/a n/a ecosystem change, USFWS (Red shiner) resources, alters indigenous fish disease vector communities, tapeworm host competes with indigenous fish Cyprinella whipplei inadequate little/no little/no competition (especially Cyprinids) for food and USFWS (Steelcolor shiner) research evidence evidence resources

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous fish for food and resources, alteration of indigenous Cyprinus carpio competition, n/a n/a n/a habitat and food webs (consumes large USFWS (Common carp, koi) ecosystem change amounts of macrophytes), increases turbidity (with feeding behaviour) Fundulus diaphanus competes with indigenous fish for food (Eastern banded n/a n/a n/a competition n/a and resources killifish) outcompetes indigenous fish for food Gambusia affinis competition, and resources (aggressive predator inadequate little/no inadequate (Western predation, behaviour), alters indigenous USFWS research evidence research mosquitofish) ecosystem change communities, alteration of food webs and habitat outcompetes indigenous fish for food Gambusia holbrooki competition, and resources (aggressive predator (Eastern n/a n/a n/a predation, behaviour), alters indigenous USFWS mosquitofish) ecosystem change communities, alteration of food webs and habitat, increases mosquitoes Gasterosteus aculeatus aculeatus little/no competes with indigenous fish for food moderate moderate competition n/a (Threespine evidence and resources stickleback) competes with indigenous fish for food Gymnocephalus inadequate little/no competition, and resources, preys on indigenous cernuus moderate USFWS research evidence predation fish, changes indigenous fish (Ruffe) communities Hypomesus competes with indigenous fish nipponensis competition, n/a n/a n/a (including salmon and shad) for food USFWS (Wakasagi smelt, disease vector and resources, parasite host Japanese smelt)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment Hypophthalmichthys harmandi competes with indigenous fish for food DFO, n/a n/a n/a competition (Largescale silver and resources USFWS carp) competes with indigenous fish for food and resources, food web changes, Hypophthalmichthys competition, changes indigenous fish communities, DFO, molitrix high high high ecosystem change, boating hazard (jumping startle USFWS (Silver carp) nuisance response), increases turbidity (with feeding behaviour) competes with indigenous fish for food and resources, food web changes, Hypophthalmichthys competition, changes indigenous fish communities, DFO, nobilis high high high ecosystem change, boating hazard (jumping startle USFWS (Bighead carp) nuisance response), increases turbidity (with feeding behaviour) Hyrcanogobius bergi n/a n/a n/a unknown unknown n/a (Volga dwarf goby) competes with indigenous fish for food competition, and resources, preys on indigenous Ictalurus furcatus little/no moderate high predation, increase fish, changes indigenous fish USFWS (Blue catfish) evidence contaminant flow communities, increases contaminant flow to avian predators Knipowitschia no evidence that species outcompetes caucasica inadequate little/no little/no not likely indigenous species for food and USFWS (Caucasian dwarf research evidence evidence resources goby) competes with indigenous fish for food Lates calcarifer competition, and resources, preys on indigenous OMNRF, n/a n/a n/a (Barramundi) predation fish, changes indigenous fish USFWS communities

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment no evidence that species outcompetes Leuciscus idus inadequate little/no disease vector, not high indigenous species for food and USFWS (Ide, orfe) research evidence likely resources, viral and parasitic host competes with indigenous fish for Leuciscus leuciscus high high moderate competition resources (including salmonid breeding USFWS (Eurasian dace) habitat) Misgurnus anguillicaudatus competition, competes with indigenous fish for food inadequate little/no little/no MNRF, (Japanese predation, disease and resources, preys on indigenous research evidence evidence USFWS weatherfish, weather vector fish, viral and parasitic host loach) unknown, competes with indigenous Monopterus albus competition, n/a n/a n/a fish for food and resources, preys on USFWS (Asian swamp eel) predation, unknown indigenous fish competes with indigenous fish for food competition, and resources, preys on indigenous fish Morone americana predation, eggs and fish, changes indigenous high moderate high USFWS (White perch) ecosystem change, zooplankton and fish communities/food hybridization webs, hybridizes with indigenous M. chrysops (white bass) competition with indigenous species for competition, food and resources, predation on Morone chrysops x predation, indigenous fish species, decline in M. saxatilis n/a n/a n/a indigenous species n/a indigenous fish species, backcrossing (Wiper) decline, risk with indigenous species (white hybridization bass) competition, competition with indigenous species for predation, food and resources, predation on Morone saxatilis n/a n/a n/a indigenous species indigenous fish species, decline in MNRF (Striped bass) decline, disease indigenous fish species, fish parasite vector host

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous fish for food competition, Mylopharyngodon and resources, preys on indigenous predation, piceus n/a n/a n/a molluscs, changes indigenous food USFWS ecosystem change, (Black carp) webs, human parasite host, fish viral disease vector and parasite host Myxocyprinus asiaticus n/a n/a n/a unknown unknown n/a (Chinese hi-fin banded shark) Neogobius fluviatilis little/no competition, competes with indigenous fish for food moderate moderate USFWS (Monkey goby) evidence disease vector and resources, fish parasite host outcompetes indigenous fish for food and resources (including spawning Neogobius competition, sites), preys on indigenous fish eggs little/no melanostomus high high ecosystem change, and fish, changes indigenous USFWS evidence (Round goby) disease vector macroinvertebrate and fish community structure and food webs, fish parasite host Odontesthes bonariensis unknown, possible indigenous fish n/a n/a n/a unknown USFWS (Argentinian community effects silverside) Oncorhynchus keta little/no competes with indigenous fish for food moderate high competition n/a (Chum salmon) evidence and resources outcompetes indigenous fish for food and resources (aggressive), changes competition, Oreochromis aureus indigenous fish community structure, n/a n/a n/a ecosystem change, n/a (Blue tilapia) restructure food web with high disease vector herbivorous diet, decreases mussel populations, fish parasite host

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment outcompetes indigenous fish for food Oreochromis competition, and resources, changes indigenous fish niloticus n/a n/a n/a USFWS ecosystem change community structure, restructures food (Nile tilapia) web with high herbivorous diet competes with indigenous fish for food competition, Osmerus mordax inadequate and resources, fish egg predator, high high ecosystem change, n/a (Rainbow smelt) research changes in indigenous fish disease vector communities, fish parasite host Parachanna africana unknown, likely predation on indigenous DFO, n/a n/a n/a unknown (Niger snakehead) fish communities USFWS competes with indigenous fish for food competition, Perca fluviatilis and resources, indigenous fish high moderate high predation, disease USFWS (Eurasian perch) predation, changes in indigenous fish vector communities, fish viral host outcompetes indigenous fish for food Perccottus glenii competition, and resources, indigenous fish high low moderate USFWS (Amur sleeper) predation predation, changes indigenous fish community structure, competition, predation/parasitism on indigenous predation, species, indirect secondary infections Petromyzon marinus little/no ecosystem change, high high from parasitism, fish parasite host, n/a (Sea lamprey) evidence indigenous species changes to indigenous fish decline, disease communities, food web changes vector competes with indigenous fish for food competition, Phoxinus phoxinus and resources, changes in indigenous moderate low low indigenous species USFWS (Eurasian minnow) fish and benthic macroinvertebrate decline communities

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous fish competition, habitat (Cyprinids) for food and resources, alteration, declines in indigenous fish species, Poecilia reticulata ecosystem change, n/a n/a n/a changes in invertebrate communities n/a (Guppy) indigenous species (predation on damselflies excludes decline, disease species), may interfere with indigenous vector fish spawning, fish parasite host Ponticola gorlap (Caspian bighead n/a n/a n/a unknown unknown n/a goby) Proterorhinus semilunaris inadequate little/no little/no competition, unknown, competes with indigenous USFWS (Freshwater research evidence evidence unknown fish for food and resources tubenose goby) competes with indigenous fish for food competition, Pseudorasbora and resources, fish parasite host little/no indigenous species OMNRF, parva high high (including the rosette agent), declines in evidence decline, disease USFWS (Stone moroko) indigenous fish species, food web vector changes competes with indigenous fish for food and resources, feeding behaviour Pterygoplichthys competition, habitat uproots aquatic macrophytes, burrowing disjunctivus alteration, n/a n/a n/a behaviour increases siltation and USFWS (Vermiculated sailfin ecosystem change, destabilizes shoreline or river banks, catfish) nuisance change nutrient flow (act as a nutrient sink)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous fish for food and resources, feeding behaviour Pterygoplichthys competition, habitat uproots aquatic macrophytes, burrowing pardalis alteration, n/a n/a n/a behaviour increases siltation and USFWS (Amazon sailfin ecosystem change, destabilizes shoreline or river banks, catfish) nuisance change nutrient flow (act as a nutrient sink) competition, unknown, competes with indigenous Rhodeus ocellatus n/a n/a n/a indigenous species fish for food and resources, decreases USFWS (Rosy bitterling) decline, unknown indigenous fish species Rhodeus sericeus unknown, not likely to have negative n/a n/a n/a not likely, unknown USFWS (Amur bitterling) impact if established competes with indigenous fish for food competition, habitat and resources, decreases indigenous alteration, Rutilus rutilus fish species, hybridization, alteration of high moderate moderate ecosystem change, USFWS (Common roach) nutrient cycle, may be able to hybridize disease vector, with indigenous fishes, fish parasite hybridization host competition, competition with indigenous fish for food predation, and resources, predation on indigenous Sander lucioperca inadequate ecosystem change, fish, declines in indigenous fish species, OMNRF, high high (Zander, pikeperch) research indigenous species food web changes, fish disease and USFWS decline, disease parasite host, possible hybridization risk vector, hybridization with indigenous species competition with indigenous fish for food Scardinius competition, habitat and resources, hybridization with inadequate inadequate erythrophthalmus moderate alteration, indigenous species (golden shiner), USFWS research research (Rudd) hybridization declines in indigenous aquatic plant species

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, competition with indigenous fish for food Silurus glanis predation, and resources, predation on indigenous little/no OMNRF, (Sheatfish, Wels high high indigenous species fish species (dominant), declines in evidence USFWS catfish) decline, disease indigenous fish species, fish virus and vector parasite host Syngnathus abaster (Black-striped inadequate little/no little/no pipefish, unknown unknown, insufficient evidence n/a research evidence evidence shortsnouted pipefish) unknown, competition with indigenous competition, habitat species for food and resources, Tinca tinca alteration, increases water turbidity with feeding n/a n/a n/a USFWS (Tench) ecosystem change, behaviour (to a lesser degree than unknown carp), changes in food webs (gastropod predator)

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Appendix C. Current and potential invasive aquatic invertebrates

Table C1. Current and potential invasive aquatic invertebrate species to the Great Lakes region, listed alphabetically by scientific name. Taxonomic family and ITIS taxonomic serial number (TSN) are listed for clarity. n/a=data not available. ITIS Scientific name Common name(s) Organism category Basis for listing TSN Anguillicoloides Eel swim bladder parasite n/a Aguilera 2015 (Anguillicola) crassus nematode Astacus astacus European crayfish crustacean, decapod 97333 CABI: invasive in Europe; Aguilera 2015 European freshwater, Austropotamobius pallipes crustacean, decapod n/a Aguilera 2015 white-clawed crayfish Bithynia tentaculata Faucet snail mollusc, gastropod 70794 Present in GLR (GLANSIS current) Brachionus leydigii Wheel animal rotifer 58475 GLANSIS watch list Bythotrephes longimanus Spiny water flea crustacean, cladoceran 684624 Present, regulated in GLR Calanipeda aquaedulcis Calanoid copepod crustacean, calanoid n/a GLANSIS watch list Cercopagis pengoi Fish hook water flea crustacean, cladoceran 684625 Present, regulated in GLR Chelicorophium Caspian mud shrimp crustacean, amphipod n/a GLANSIS watch list; Pagnucco et al. 2015 curvispinum Cherax destructor Yabbie, Yabby crustacean, decapod n/a Regulated in GLR Cherax tenuimanus Marron crustacean, decapod n/a Regulated in GLR Chinese mystery Cipangopaludina chinensis mollusc, gastropod 70329 Present, regulated in GLR snail Japanese trapdoor Cipangopaludina japonica mollusc, gastropod 70332 Present in GLR (GLANSIS current) snail, mystery snail Corbicula fluminea Asian clam mollusc, bivalve 81387 Present, regulated in GLR Cordylophora caspia Freshwater hydroid cnidarian, hydrozoan 48893 Present in GLR (GLANSIS current) Cornigerius maeoticus Water flea crustacean, cladoceran n/a Present in GLR (GLANSIS current) Cyclops kolensis Cyclopoid copepod crustacean, copepod 88685 GLANSIS watch list Daphnia cristata Water flea crustacean, cladoceran n/a GLANSIS watch list

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ITIS Scientific name Common name(s) Organism category Basis for listing TSN Daphnia lumholtzi Water flea crustacean, cladoceran 684652 Present, regulated in GLR Dikerogammarus Scud, demon shrimp crustacean, amphipod n/a GLANSIS watch list haemobaphes Dikerogammarus villosus Killer shrimp crustacean, amphipod n/a GLANSIS; Pagnucco et al. 2015 Dreissena polymorpha Zebra mussel mollusc, bivalve 81339 Regulated in GLR Dreissena rostriformis Quagga mussel mollusc, bivalve 567514 Regulated in GLR bugensis Echinogammarus ischnus Scud crustacean, amphipod 656834 Present in GLR (GLANSIS current) Echinogammarus trichiatus Scud crustacean, amphipod n/a USGS: present in GLR; Aguilera 2015 Echinogammarus Scud crustacean, amphipod n/a GLANSIS watch list warpachowskyi Harpacticoid Ectinosoma abrau crustacean, copepod n/a GLANSIS watch list copepod Eriocheir sinensis Chinese mitten crab crustacean, decapod 99058 Regulated in GLR, Watch list in GLR Faxonius (Orconectes) Spinycheek crayfish crustacean, decapod 97423 Aguilera 2015 limosus Faxonius (Orconectes) Rusty crayfish crustacean, decapod 97424 Regulated in GLR, Watch list in GLR rusticus Filinia cornuta Wheel animal rotifer n/a GLANSIS watch list Filinia passa Wheel animal rotifer 59430 GLANSIS watch list Moss animal, Fredericella sultana bryozoan 156723 GLANSIS watch list branching bryozoan Gammarus tigrinus Amphipod crustacean, amphipod 93781 Present in GLR (GLANSIS current) Gmelinoides fasciatus Baikalian amphipod crustacean, amphipod n/a Pagnucco et al. 2015 Gyrodactylus salaris Salmon fluke parasite 54834 Aguilera 2015; CABI: invasive in northern Europe Hemimysis anomala Bloody shrimp crustacean, mysid 90568 Present in GLR (GLANSIS current) Heterocope appendiculata Calanoid copepod crustacean, copepod 550401 GLANSIS watch list Heterocope caspia Calanoid copepod crustacean, copepod n/a GLANSIS watch list Hypania invalida Bristle worm polychaete 67824 Aguilera 2015; Straka et al. 2015

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ITIS Scientific name Common name(s) Organism category Basis for listing TSN Hypanis (Monodacna) Clam, coloured mollusc, bivalve n/a GLANSIS watch list colorata lagoon cockle crustacean, copepod, Lernaea cyprinacea Anchor worm 89254 CABI: invasive worldwide, Aguilera 2015 parasite Trematode flatworm, Leyogonimus polyoon trematode, parasite n/a GLANSIS watch list leo Opossum shrimp, Limnomysis benedeni Caspian slender crustacean, mysid 90553 CABI: invasive in Europe; Aguilera 2015 shrimp Limnoperna fortunei Golden mussel mollusc, bivalve n/a USFWS Malaysian trumpet CABI: invasive worldwide, present in Southern Melanoides tuberculatus snail, red-rimmed mollusc, gastropod 71533 US; Aguilera 2015 melania Obesogammarus crassus Scud, thick scud crustacean, amphipod n/a GLANSIS watch list Obesogammarus obesus Scud crustacean, amphipod n/a GLANSIS watch list Paraleptastacus Harpacticoid crustacean, copepod 86909 GLANSIS watch list spinicauda triseta copepod Paramysis (Mesomysis) Opossum shrimp crustacean, mysid 90314 GLANSIS watch list intermedia Paramysis (Metamysis) Opossum shrimp crustacean, mysid 90324 GLANSIS watch list ullskyi Paramysis (Serrapalpisis) Opossum shrimp crustacean, mysid 90318 GLANSIS watch list lacustris Podonevadne trigona Water flea crustacean, cladoceran n/a USFWS; Aguilera 2015 ovum Sturgeon egg Polypodium hydriforme parasite 51255 Aguilera 2015 parasite Pomacea canaliculata Applesnail mollusc, gastropod 568121 CABI: invasive worldwide; Aguilera 2015 Pontogammarus Scud crustacean, amphipod n/a GLANSIS watch list; Aguilera 2015 robustoides Potamopyrgus New Zealand mollusc, gastropod 205006 Regulated in GLR antipodarum mudsnail

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ITIS Scientific name Common name(s) Organism category Basis for listing TSN Procambarus clarkii Red swamp crayfish crustacean, decapod 97491 Present in GLR (GLANSIS current) Procambarus fallax f. Marmorkrebs, crustacean, decapod 97514 USFWS virginalis marbled crayfish Pseudodactylogyrus Eel parasite parasite n/a Aguilera 2015 anguillae Pseudodactylogyrus bini Eel gill parasite parasite n/a Aguilera 2015 Radix auricularia European ear snail mollusc, gastropod 76526 Present in GLR (GLANSIS current) Schyzocotyle Asian fish tapeworm, CABI: invasive worldwide; detected in GLR, (Bothriocephalus) parasite 1059846 Asian tapeworm Marcogliese et al. 2016 acheilognathi Sinanodonta (Anodonta) Chinese pond mussel mollusc, bivalve 983676 Regulated in GLR woodiana Valvata piscinalis European valve snail mollusc, gastropod 70348 Present in GLR (GLANSIS current) Viviparus georgianus Banded mystery snail mollusc, gastropod 70307 Present in GLR (GLANSIS current)

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Table C2. Geographic ranges and pathways for current and potential invasive aquatic invertebrates to the Great Lakes region (GLR), listed alphabetically by scientific name. Origin (indigenous geographic range), basis for inclusion in list, GLR restrictions, establishment, current ranges, and pathways/vectors of initial entry listed for each species. n/a=data not available or applicable, italics indicate potential pathways for species yet to arrive in the GLR. GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Anguillicoloides aquarium/garden, North commercial crassus Date NY (Hudson hitchhiker, live Asia Eastern Asia n/a America, shipping, canal, (Eel swim-bladder unknown River) trade, recreational Europe bait release nematode) boating commercial Astacus astacus shipping, canal, (European Europe n/a n/a n/a n/a Europe aquarium/garden recreational crayfish) boating, bait release commercial Austropotamobiu shipping, canal, s pallipes Europe n/a n/a n/a n/a Europe unknown recreational (White-clawed boating, bait crayfish) release Great Lakes Bithynia North recreational (Erie, Ontario, commercial tentaculata Europe n/a n/a 1871 America, boating, canal, Michigan and shipping (Faucet snail) Europe bait release their drainages) Africa, Northern Africa, Brachionus recreational Oceania, Africa, Oceania, commercial leydigii n/a n/a n/a boating, canal, Asia, Australia, Asia, shipping (Wheel animal) bait release Europe Asia, Europe Europe Bythotrephes Great Lakes North recreational Northern commercial longimanus Eurasia IL, MN 1984 (all), throughout America, boating, canal, Europe, Asia shipping (Spiny waterflea) GLR Eurasia bait release

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Calanipeda Africa, Africa, Africa, recreational aquaedulcis commercial Europe, Europe, n/a n/a n/a Europe, boating, canal, (Calanoid shipping Eurasia Eurasia Eurasia bait release copepod) Cercopagis Ponto- Great Lakes North recreational pengoi commercial Eurasia Caspian IL 1998 (Erie, Michigan, America, boating, canal, (Fish hook shipping basin Ontario), NY Eurasia bait release waterflea) Chelicorophium Ponto- recreational curvispinum commercial Eurasia Caspian n/a n/a n/a Europe boating, canal, (Caspian mud shipping basin bait release shrimp) commercial shipping, canal, Cherax destructor IL, MN, Oceania Australia n/a n/a Europe aquarium/garden recreational (Yabbie, yabby) OH, ON boating, bait release commercial Cherax shipping, canal, Africa, tenuimanus Australia Australia IL, OH n/a n/a aquarium/garden recreational Oceania (Marron) boating, bait release Great Lakes (Erie, upper St. Cipangopaludina Lawrence), recreational chinensis Southeast North aquarium/garden, Asia IL, MN 1931 Lake Ontario boating, canal, (Chinese mystery Asia America live trade and Michigan bait release snail) basins, ON, MI, IN, OH, WI, NY

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range commercial Cipangopaludina shipping, Great Lakes japonica Japan, recreational (Erie, North (Japanese Asia Taiwan, n/a 1940 intentional release boating, canal, Michigan), OH, America trapdoor, mystery Korea aquarium/garden, MI, WI, PA snail) live trade, bait release North commercial Great Lakes America, Corbicula shipping, (Erie, Michigan, South aquarium/garden, fluminea Asia n/a IL, IN 1980 recreational Superior), USA America, live trade (Asian clam) boating, canal, (widespread) Europe, bait release Africa Cordylophora commercial recreational caspia Date Great Lakes North Eurasia n/a n/a shipping, boating, canal, (Freshwater unknown (Erie), IL, NY America aquarium/garden bait release hydroid) Ponto- Caspian Cornigerius canal, recreational basin, commercial maeoticus Eurasia n/a n/a n/a Europe boating, bait Danube, shipping (Water flea) release Dnieper and Bug rivers Cyclops kolensis Ponto- North canal, recreational commercial (Cyclopoid Eurasia Caspian n/a n/a n/a America, boating, bait shipping copepod) basin Europe release Ponto- canal, recreational Daphnia cristata commercial Eurasia Caspian n/a n/a n/a Eurasia boating, bait (Water flea) shipping basin release Africa, East Africa, Daphnia lumholtzi Illinois River, IL, North recreational Asia, Australia, IL 1999 canal, bait release (Water flea) OH, MI, WI America boating Oceania India

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Dikerogammarus Ponto- canal, recreational haemobaphes commercial Eurasia Caspian n/a n/a n/a Europe boating, bait (Scud, demon shipping basin release shrimp) Dikerogammarus Ponto- canal, recreational commercial villosus Eurasia Caspian ON n/a n/a Europe boating, bait shipping (Killer shrimp) basin release Dreissena Ponto- IL, IN, MI, Great Lakes North recreational commercial polymorpha Eurasia Caspian MN NY, 1986 (all), throughout America, boating, canal, shipping (Zebra mussel) basin OH GLR Europe bait release Dreissena Ponto- IL, IN, MI, Great Lakes North recreational rostriformis commercial Eurasia Caspian MN NY, 1989 (all), throughout America, boating, canal, bugensis shipping basin OH GLR Europe bait release (Quagga mussel) Echinogammarus Ponto- Great Lakes Eurasia, commercial recreational ischnus Eurasia Caspian n/a 1993 (all), throughout North shipping boating, canal (Scud) basin GLR America Echinogammarus Ponto- commercial canal, recreational trichiatus Eurasia Caspian n/a n/a n/a Eurasia shipping boating (Scud) basin Echinogammarus Ponto- commercial canal, recreational warpachowskyi Eurasia Caspian n/a n/a n/a Eurasia shipping boating (Scud) basin Ectinosoma Ponto- Europe, canal, recreational abrau commercial Eurasia Caspian n/a n/a n/a North boating, bait (Harpacticoid shipping basin America release copepod) commercial Eriocheir sinensis Eurasia, canal, recreational IL, NY, shipping, (Chinese mitten Asia China, Korea n/a n/a North boating, bait PQ aquarium/garden, crab) America release live trade

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Turkey, Russia, Eurasia, Brazil, South Mexico, canal, recreational Filinia cornuta America, commercial Austria, n/a n/a n/a n/a boating, bait (Wheel animal) North shipping Hungary, release America, New Oceania Zealand, China canal, recreational Filinia passa commercial Oceania Australia n/a n/a n/a Europe boating, bait (Wheel animal) shipping release Europe, Asia, Eurasia, Fredericella Oceania, canal, recreational Oceania, North commercial sultana Western n/a n/a n/a boating, bait North America shipping (Moss animal) North release America America Gammarus Great Lakes Europe, North commercial recreational tigrinus Atlantic coast n/a 2001 (all), throughout North America shipping boating, canal (Amphipod) GLR America Gmelinoides fasciatus commercial canal, recreational Eurasia Asia, Siberia n/a n/a n/a Eurasia (Baikalian shipping boating amphipod) commercial Gyrodactylus Scandinavia, aquarium/garden, shipping, salaris Eurasia Western n/a n/a n/a Eurasia live trade, recreational (Salmon fluke) Russia hitchhiker boating, canal

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Great Lakes (Michigan, Erie, Hemimysis Ponto- Ontario, likely Eurasia, anomala commercial recreational Eurasia Caspian WI 2006 present in North (Bloody red shipping boating, canal basin Huron and America shrimp) Superior), ON, PQ, MI, NY Heterocope Ponto- canal, recreational appendiculata commercial Eurasia Caspian n/a n/a n/a Europe boating, bait (Calanoid shipping basin release copepod) Heterocope Ponto- canal, recreational caspia commercial Eurasia Caspian n/a n/a n/a Eurasia boating, bait (Calanoid shipping basin release copepod) Ponto- Hypania invalida commercial canal, recreational Eurasia Caspian n/a n/a n/a Europe (Bristle worm) shipping boating basin Hypanis colorata Ponto- recreational commercial (Clam, coloured Eurasia Caspian n/a n/a n/a Eurasia boating, canal, shipping lagoon cockle) basin bait release Africa, North commercial Lernaea Eurasia, Eurasia, America, shipping, cyprinacea cryptogeni n/a n/a n/a aquarium/garden cryptogenic South recreational (Anchor worm) c America, boating, canal Eurasia commercial Leyogonimus shipping, polyoon Eastern Mississippi North Europe n/a 2007 recreational bait release (Trematode Europe River basin, WI America boating, canal, flatworm, leo) hitchhiker

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Limnomysis benedeni Ponto- commercial canal, recreational (Opossum Eurasia Caspian n/a n/a n/a Eurasia shipping boating shrimp, Caspian basin slender shrimp) Limnoperna IL, MI, Asia, canal, recreational Southeast commercial fortunei Asia OH, ON, n/a n/a South boating, bait Asia shipping (Golden mussel) WI America release North Melanoides America, commercial tuberculatus Northern Africa, shipping, (Malaysian Africa, Africa, n/a n/a n/a Eurasia, aquarium/garden recreational trumpet snail, Asia Southern South boating, canal, red-rimmed Asia America, bait release melania) Oceania Ponto- Obesogammarus Caspian commercial recreational crassus Eurasia n/a n/a n/a Eurasia basin, Baltic shipping boating, canal (Scud, thick scud) Sea Obesogammarus Ponto- commercial recreational obesus Eurasia Caspian n/a n/a n/a Eurasia shipping boating, canal (Scud) basin eastern commercial Faxonius limosus Atlantic North Eurasia, bait release, shipping, (Spinycheek watershed, n/a n/a n/a America Africa aquarium/garden recreational crayfish) Lake Erie boating, canal (cryptogenic) Ohio River commercial IL, MI, Great Lakes Faxonius rusticus North basin, Lake North bait release, shipping, MN, PA, 1800s (all), throughout (Rusty crayfish) America Erie America aquarium/garden recreational PQ GLR (cryptogenic) boating, canal

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Paraleptastacus Ponto- Eurasia, recreational spinicauda triseta commercial Eurasia Caspian n/a n/a n/a North boating, canal, (Harpacticoid shipping basin America bait release copepod) Paramysis Ponto- intermedia commercial recreational Eurasia Caspian n/a n/a n/a Eurasia (Opossum shipping boating, canal basin shrimp) Paramysis (Metamysis) Ponto- commercial recreational ullskyi Eurasia Caspian n/a n/a n/a Eurasia shipping boating, canal (Opossum basin shrimp) Paramysis Ponto- lacustris commercial recreational Eurasia Caspian n/a n/a n/a Eurasia (Opossum shipping boating, canal basin shrimp) Podonevadne Ponto- recreational commercial trigona ovum Eurasia Caspian n/a n/a n/a Eurasia boating, canal, shipping (Water flea) basin bait release Polypodium Ponto- commercial recreational hydriforme Eurasia Caspian n/a n/a n/a Eurasia shipping, boating, canal, (Sturgeon egg basin hitchhiker bait release parasite) North America, live trade, Pomacea Africa, recreational South Argentina, aquarium/garden, canaliculata n/a n/a n/a Eurasia, boating, canal, America Uruguay commercial (Applesnail) South bait release shipping America, Oceania

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Pontogammarus Ponto- commercial recreational robustoides Eurasia Caspian n/a n/a n/a Eurasia shipping boating, canal (Scud) basin Great Lakes (Erie, Ontario, Europe, Potamopyrgus Michigan, Asia, antipodarum Superior), ON commercial recreational Oceania New Zealand MN, WI 1991 North (New Zealand (Welland Canal, shipping boating, canal America, mudsnail) Thunder Bay), Oceania OH, PA, IL, NY, MN, WI Europe, Mississippi Procambarus Great Lakes Asia, River bait release, live clarkii North (Erie, North commercial drainage, n/a 1967 trade, recreational (Red swamp America Michigan), OH, America, shipping, canal Gulf Coast boating crayfish) WI South plain America unknown, discovered in commercial Procambarus German live Europe, shipping, fallax f. virginalis North aquarium/garden, trade, no n/a n/a n/a Africa, recreational (Marmorkrebs, America live trade indigenous Asia boating, canal, marbled crayfish) population bait release reported commercial Pseudodactylogyr Europe, shipping, aquarium/garden, us anguillae Asia East Asia n/a n/a n/a North recreational live trade (Eel parasite) America boating, canal, bait release

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range commercial North Pseudodactylogyr aquarium/garden, shipping, America, us bini Asia Japan n/a n/a n/a live trade, recreational Asia, (Eel gill parasite) hitchhiker boating, canal, Europe bait release Great Lakes (Huron), local populations in Lake Ontario, North commercial Radix auricularia Europe, Erie and America, shipping, (European ear Eurasia n/a 1869 aquarium/garden North Asia Michigan Oceania, recreational snail) drainages, NY Eurasia boating, canal (Finger Lakes), VT (Lake Champlain) North America, commercial Schyzocotyle Africa, shipping, canal, aquarium/garden, acheilognathi Asia Eastern Asia n/a n/a n/a Eurasia, recreational hitchhiker (Asian tapeworm) South boating, bait America, release Oceania North commercial Sinanodonta America, shipping, canal, woodiana Asia Eastern Asia IL n/a n/a Europe, aquarium/garden recreational (Chinese pond South boating, bait mussel) America release

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GLR Current Species GLR Current GLR Introduction GLR spread Origin Origin detail regulated invaded (common name) year range pathway pathway locations range Great Lakes (Erie, Ontario, Europe, Superior, MI), commercial Valvata piscinalis Caucasus, St. Lawrence North hitchhiker, shipping, (European valve Eurasia n/a 1897 Siberia, River, NY America aquarium/garden recreational snail) Central Asia (Finger Lakes), boating, canal VT (Lake Champlain) Mississippi and St. unauthorized Viviparus Lawrence Lake Erie, commercial intentional georgianus North River Ontario and North shipping, n/a 1867 release, (Banded mystery America systems, Michigan America recreational aquarium/garden, snail) Northeastern drainages boating, canal live trade USA to Gulf of Mexico

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Table C3. Size and habitat preferences for current and potential invasive aquatic invertebrates to the Great Lakes region, listed alphabetically by scientific name. n/a=data not available. Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight Anguillicoloides crassus freshwater, ectoparasitic, completes life cycle through eel (Eel swim-bladder 3 cm n/a 2–12 months brackish, fish parasite and intermediate hosts nematode) marine Astacus astacus lake/pond, benthic, littoral vegetation, soft substrates 16 cm n/a 20 years freshwater (European crayfish) river/stream (sand) Austropotamobius lake/pond, benthic, submerged pallipes 12 cm 90 g 10 years freshwater river/stream rocks/cobbles/logs/roots/leaves (White-clawed crayfish) Bithynia tentaculata 17–39 lake/pond, littoral, benthic: on substrates in fall/winter, on 15 mm n/a freshwater (Faucet snail) months river/stream aquatic macrophytes in spring/summer Brachionus leydigii lake/pond, littoral zooplankton: mainly among aquatic 280 µm n/a 6–45 days freshwater (Wheel animal) river/stream vegetation, thrive in eutrophic conditions Bythotrephes longimanus pelagic zooplankton, oligotrophic and 15 mm n/a 12–26 days freshwater lake/pond (Spiny waterflea) mesotrophic lakes Calanipeda aquaedulcis 120–1900 freshwater, pelagic and littoral zooplankton, oligotrophic n/a n/a lake/pond (Calanoid copepod) µm brackish and mesotrophic lakes Cercopagis pengoi 6–13 mm freshwater, pelagic zooplankton, oligotrophic and n/a 3–15 days lake/pond (Fish hook waterflea) (with tail) brackish mesotrophic lakes Chelicorophium freshwater, littoral, benthic: mud tubes on hard substrate lake/pond, curvispinum 10 mm n/a 8-12 months brackish, (rock, wood, submerged vegetation, bivalve river/stream (Caspian mud shrimp) marine shells) lake/pond, Cherax destructor littoral, benthic: muddy substrates with 30 cm 300 g 3–7 years freshwater river/stream, (Yabbie, yabby) moderate turbidity, dig borrows 0.5 - 2m deep wetland littoral, benthic: silty, sandy substrates with Cherax tenuimanus freshwater, 40 cm 2.5 kg 5–10 years river/stream high organic matter and refugia (rocks, tree (Marron) brackish roots), require good water quality

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Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight Cipangopaludina lake/pond, littoral, benthic, lentic or slow-moving water, chinensis 65 mm n/a 3–5 years freshwater river/stream, soft substrate, silt, sand, mud (Chinese mystery snail) wetland littoral, epibenthic, endobenthic with soft Cipangopaludina japonica lake/pond, substrate (mud, sand, vegetation, wood), (Japanese trapdoor, 50 mm n/a 8 years freshwater river/stream, lentic or slow-moving water, meso and mystery snail) wetland eutrophic lakes, coarse woody debris, unstructured habitats, grass beds littoral, epibenthic, endobenthic with soft Corbicula fluminea freshwater, lake/pond, substrate (sand, gravel, and also mud, silt), 50 mm n/a 1–7 years (Asian clam) brackish river/stream requires high dissolved oxygen and prefers moving water, intolerant of pollution 5 cm lake/pond, Cordylophora caspia (colony), 1 freshwater, benthic: attached to hard surfaces (rocks, n/a n/a river/stream, (Freshwater hydroid) mm brackish pilings, vegetation, wood, shells) wetland (polyp) Cornigerius maeoticus freshwater, lake/pond, pelagic zooplankton, prefers metalimnion, 0.5–1 mm n/a n/a (Water flea) brackish river/stream standing or slow flowing water Cyclops kolensis pelagic, littoral zooplankton, thrives in 1.5 mm n/a 1–2 months freshwater lake/pond (Cyclopoid copepod) eutrophic water Daphnia cristata 0.6 mm n/a 1-2 months freshwater lake/pond pelagic, prefers eutrophic water (Water flea) Daphnia lumholtzi 3.5 mm n/a 1 month freshwater lake/pond pelagic (Water flea) Dikerogammarus lake/pond, epibenthic, epiphytic: solid substrates, freshwater, haemobaphes 21 mm n/a 1 year river/stream, including live zebra mussel shells, brackish (Scud, demon shrimp) wetland macrophytes, filamentous algae epibenthic, epiphytic: all solid substrates lake/pond, Dikerogammarus villosus freshwater, including zebra mussel beds, rocks, artificial 30 mm n/a 1 year river/stream, (Killer shrimp) brackish surfaces, macrophytes), prefers slow-moving wetland water

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Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight epibenthic, epiphytic (attaches to most freshwater, lake/pond, Dreissena polymorpha substrates): rock, macrophytes, artificial 50 mm n/a 3-9 years brackish, river/stream, (Zebra mussel) surfaces, macroinvertebrates, will also attach marine wetland to soft substrate habitats Dreissena rostriformis freshwater, lake/pond, epibenthic: both hard and soft substrates, bugensis 4 cm n/a 4-5 years brackish river/stream including sand and mud (Quagga mussel) benthic, wide habitat and substrate range, prefers shallow margins of moving water with Echinogammarus ischnus freshwater, lake/pond, 11 mm n/a 1 year hard substrates (gravel, rock, mussels), (Scud) brackish river/stream tolerates soft substrates, closely associated with zebra mussel colonies Echinogammarus freshwater, lake/pond, trichiatus 10 mm n/a n/a benthic brackish river/stream (Scud) Echinogammarus freshwater, lake/pond, benthic, prefers meso and eutrophic, well warpachowskyi 7 mm n/a 1 year brackish river/stream oxygenated water (Scud) freshwater, Ectinosoma abrau lake/pond, >0.6 mm n/a n/a brackish, pelagic (Harpacticoid copepod) river/stream marine 10 cm freshwater, lake/pond, Eriocheir sinensis littoral, benthic, slow or still water, repro in salt (carapace n/a 1–5 years brackish, river/stream, (Chinese mitten crab) water width) marine wetland benthic, wide range of habitats, fast and still Faxonius limosus lake/pond, water, prefers cryptic habitat for shelter with 11 cm n/a 2–4 years freshwater (Spinycheek crayfish) river/stream clear, well oxygenated, slow-flowing water, soft substrates allow species to burrow benthic, wide range of habitats, fast and still Faxonius rusticus lake/pond, water, prefers cryptic habitat for shelter with 10 cm n/a 2–4 years freshwater (Rusty crayfish) river/stream clear, well oxygenated water, soft substrates allow species to burrow

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Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight Filinia cornuta pelagic, associated with eutrophication and <200 µm n/a 6–45 days freshwater lake/pond (Wheel animal) organic pollution Filinia passa lake/pond, <200 µm n/a 6–45 days freshwater pelagic, tolerates eutrophication (Wheel animal) river/stream 1.5 mm (individual 1–20 years epibenthic, epiphytic: solid substrates, Fredericella sultana zooid), (theoretically lake/pond, including underside of stones, leaves and n/a freshwater (Moss animal) colonies of indefinitely river/stream roots, prefers near-shore, flowing nutrient rich 100s of as a colony) water zooids benthic, tolerates eutrophic, hypoxic Gammarus tigrinus freshwater, lake/pond, conditions, prefers rivers and shallow lakes 11 mm n/a 2 months (Amphipod) brackish river/stream with turbid water, soft substrates and vegetation Gmelinoides fasciatus freshwater, lake/pond, 14 mm n/a 1 year benthic (Baikalian amphipod) brackish river/stream ectoparasitic, skin and gills of Atlantic and Gyrodactylus salaris freshwater, lake/pond, Baltic salmon, also can survive and reproduce 1 mm n/a 6–15 days (Salmon fluke) brackish river/stream on other indigenous salmonids, completes life cycle with fish host pelagic, profundal zone, prefers slow or still Hemimysis anomala freshwater, lake/pond, 13 mm n/a 9 months water with hard substrate with shelter and (Bloody red shrimp) brackish river/stream scarce vegetation Heterocope appendiculata lake/pond, pelagic, littoral zooplankton, thrives in 2.2 mm n/a 20–30 days freshwater (Calanoid copepod) river/stream eutrophic water Heterocope caspia freshwater, 2 mm n/a n/a lake/pond pelagic, above thermocline (Calanoid copepod) brackish benthic, littoral, lives within muddy tube on Hypania invalida freshwater, lake/pond, various substrates, prefers soft substrates 33.5 mm n/a 10 months (Bristle worm) brackish river/stream (mud, gravel, silt, clay, fine sand) and slow to still water

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Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight Hypanis colorata freshwater, lake/pond, benthic, prefers shallow, oxygen rich water (Clam, coloured lagoon 41 mm n/a 2–5 years brackish river/stream with soft substrates wit cockle) lake/pond, pelagic, ectoparasitic, requires fish host Lernaea cyprinacea 11 mm n/a 1–3 months freshwater river/stream, (usually cyprinid, gills and skin) to complete (Anchor worm) fish parasite life cycle, no intermediate host lake/pond, endoparasitic, requires two intermediate hosts river/stream, Leyogonimus polyoon (including faucet snails and other benthic 1 mm n/a n/a freshwater benthic (Trematode flatworm, leo) macroinvertebrates) and one definitive host invertebrate/ (waterfowl, duodenum and intestine) bird parasite Limnomysis benedeni wide habitat and substrate range, prefers freshwater, lake/pond, (Opossum shrimp, 13 mm n/a n/a shallow, littoral still to slow water with cryptic brackish river/stream Caspian slender shrimp) habitat Limnoperna fortunei freshwater, lake/pond, 46 mm n/a 2–10 years epibenthic, requires hard substrates (Golden mussel) brackish river/stream Melanoides tuberculatus freshwater, lake/pond, benthic, wide habitat and substrate range, (Malaysian trumpet snail, 80 mm n/a 2–3 years brackish, river/stream, prefers permanent water with emergent plants red-rimmed melania) marine wetland and soft substrates Obesogammarus crassus freshwater, lake/pond, benthic, tolerates wide range of habitats and 12 mm n/a n/a (Scud, thick scud) brackish river/stream water quality Obesogammarus obesus freshwater, lake/pond, benthic, wide range of habitats, prefers slow, 11 mm n/a <1 year (Scud) brackish river/stream deep water Paraleptastacus freshwater, lake/pond, benthic, inhabits soft sediments (sand, spinicauda triseta 2 mm n/a 1–6 months brackish, river/stream beaches), well oxygenated water (Harpacticoid copepod) marine Paramysis intermedia freshwater, lake/pond, benthic, prefers shallow slow or still water 13 mm n/a <1 year (Opossum shrimp) brackish river/stream with sandy substrates Paramysis (Metamysis) freshwater, lake/pond, benthic, prefers shallow, sandy substrates ullskyi 13 mm n/a <1 year brackish river/stream with slow to still water (Opossum shrimp)

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Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight benthic, tolerates wide range of habitats and Paramysis lacustris freshwater, lake/pond, 17 mm n/a <1 year water quality, thrives in eutrophic water with (Opossum shrimp) brackish river/stream soft substrates (sand, silt) Podonevadne trigona pelagic, epibenthic, still to slow-moving water, freshwater, lake/pond, ovum 0.5 mm n/a 1–2 months submerged and emergent macrophytes, brackish river/stream (Water flea) cryptic environments endoparasitic, acipenseriform fishes Polypodium hydriforme lake/pond, 4 mm n/a 1–10 years freshwater (sturgeons and paddlefish, gonads) are the (Sturgeon egg parasite) river/stream definitive host littoral, prefers still water with vegetation and lake/pond, Pomacea canaliculata soft substrates (mud), facilitative air breather, 7.5 cm n/a 2–5 years freshwater river/stream, (Applesnail) will leave water at night to feed on fresh wetland vegetation Pontogammarus freshwater, lake/pond, benthic, prefers still to slow-moving water with robustoides 20 mm n/a 2–6 months brackish river/stream vegetation, substrates of stone, sand or mud (Scud) littoral, tolerates wide range of habitats, Potamopyrgus lake/pond, freshwater, turbidity, fast to still water, thrives in eutrophic antipodarum 6 mm n/a 1–2 years river/stream, brackish water, able to burrow into soft substrates, (New Zealand mudsnail) wetland early successional species tolerates a wide range of habitats, fast to still lake/pond, Procambarus clarkii water, prefers soft substrates (mud, sand) 12 cm n/a - 2–5 years freshwater river/stream, (Red swamp crayfish) with organic debris and cryptic habitat, able to wetland create burrows in fine sediments tolerates a wide range of habitats, prefers soft Procambarus fallax f. lake/pond, substrates (mud, sand) with organic debris, virginalis 13 cm n/a 2–4 years freshwater river/stream, cryptic habitat and slow to still water, burrows (Marmorkrebs, Marbled wetland only under extreme conditions (tertiary crayfish) burrower) Pseudodactylogyrus freshwater, lake/pond, anguillae 1.5 mm n/a 2–4 months brackish, ectoparasitic, found on gills of eels river/stream (Eel parasite) marine

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Maximum Maximum Species Salinity Habitat adult adult Longevity Habitat description (common name) tolerance preference length weight freshwater, Pseudodactylogyrus bini lake/pond, 2 mm n/a 2–4 months brackish, ectoparasitic, found on gills of eels (Eel gill parasite) river/stream marine lake/pond, benthic, wide habitat and substrate range, Radix auricularia freshwater, 35 mm n/a 1 year river/stream, prefers still to slow eutrophic water with soft (European ear snail) brackish wetland substrates (sand, silt) Schyzocotyle completes life cycle through a variety of fish acheilognathi 8–100 cm n/a n/a freshwater fish parasite (particularly cyprinids) and copepod hosts (Asian tapeworm) benthic, silt, clay substrates, turbid conditions, Sinanodonta woodiana lake/pond, 30 cm n/a 12–14 years freshwater standing or slow flowing water, indiscriminant (Chinese pond mussel) river/stream obligate ectoparasite larval stage on fish lake/pond, benthic, wide habitat and substrate range, Valvata piscinalis 7 mm n/a 1–2 years freshwater river/stream, prefers still to slow clear water with soft (European valve snail) wetland substrates (sand, mud, silt) lake/pond, benthic, wide habitat and substrate range, Viviparus georgianus 45 mm n/a 1–3 years freshwater river/stream, prefers still to slow eutrophic water with soft (Banded mystery snail) wetland substrates (sand, mud, silt)

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Table C4. Diet and reproductive behaviour for current and potential invasive aquatic invertebrates to the Great Lakes region, listed alphabetically by scientific name. n/a=data not available. Species Age to Trophic guild Diet details Reproduction (common name) maturity Anguillicoloides crassus ectoparasite fish parasite 1 month sexual, asexual (Eel swim-bladder nematode) Astacus astacus omnivore benthic invertebrates, plants 4 years sexual (European crayfish) Austropotamobius pallipes omnivore benthic invertebrates, plants 3–4 years sexual (White-clawed crayfish) Bithynia tentaculata omnivore algae, benthic macroinvertebrates 6 months sexual (Faucet snail) Brachionus leydigii herbivore bacteria, detritus 1 day sexual, asexual (Wheel animal) Bythotrephes longimanus carnivore zooplankton 5–13 days sexual, asexual (Spiny waterflea) Calanipeda aquaedulcis omnivore zooplankton, phytoplankton, detritus n/a sexual, asexual (Calanoid copepod) Cercopagis pengoi carnivore small zooplankton 15 days sexual, asexual (Fish hook waterflea) Chelicorophium curvispinum herbivore phytoplankton, detritus 3 months sexual (Caspian mud shrimp) Cherax destructor mainly rotting vegetation and other detritus, but also omnivore 1 year sexual (Yabbie, yabby) fish, plants, benthic macroinvertebrates, zooplankton Cherax tenuimanus mainly rotting vegetation and other detritus, but also omnivore 1–3 years sexual (Marron) fish, plants, benthic macroinvertebrates Cipangopaludina chinensis organic and inorganic benthos, benthic and epiphytic omnivore 1 year sexual (Chinese mystery snail) algae, diatoms Cipangopaludina japonica (Japanese trapdoor, mystery herbivore detritus, sludge, diatoms, periphyton, phytoplankton 1 year sexual snail) Corbicula fluminea omnivore zooplankton, phytoplankton, detritus 1 year sexual, asexual (Asian clam)

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Species Age to Trophic guild Diet details Reproduction (common name) maturity Cordylophora caspia omnivore, zooplankton, benthic macroinvertebrates, small fish n/a sexual (Freshwater hydroid) carnivore Cornigerius maeoticus carnivore zooplankton n/a sexual, asexual (Water flea) Cyclops kolensis carnivore zooplankton n/a sexual (Cyclopoid copepod) Daphnia cristata herbivore phytoplankton 5–10 days sexual, asexual (Water flea) Daphnia lumholtzi herbivore phytoplankton, organic detritus, bacteria 5–9 days sexual (Water flea) Dikerogammarus haemobaphes omnivore detritus, sediments, algae, benthic macroinvertebrates 1–2 months sexual (Scud, demon shrimp) macroinvertebrates, detritus, algae, fish eggs, small Dikerogammarus villosus omnivore fish; extremely aggressive, observed to kill potential 1 month sexual (Killer shrimp) prey without consuming it Dreissena polymorpha phytoplankton, bacteria, protozoans, microzooplankton, herbivore 1–2 years sexual (Zebra mussel) silt Dreissena rostriformis phytoplankton, bacteria, protozoans, microzooplankton, bugensis omnivore 1 year sexual silt (Quagga mussel) Echinogammarus ischnus omnivore detritus, benthic macroinvertebrates, plant material 1–2 months sexual (Scud) Echinogammarus trichiatus omnivore, detritus, benthic macroinvertebrates, plant omnivore n/a sexual (Scud) material Echinogammarus warpachowskyi omnivore detritus, benthic macroinvertebrates, plant material 1–2 months sexual (Scud) Ectinosoma abrau unknown unknown n/a sexual (Harpacticoid copepod) Eriocheir sinensis detritus, benthic macroinvertebrates, plant material, omnivore 3–5 years sexual (Chinese mitten crab) zooplankton, small fish

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Species Age to Trophic guild Diet details Reproduction (common name) maturity opportunistic, macrophytes, benthic invertebrates Faxonius limosus omnivore (including snails), detritus, periphyton, fish eggs, small 1–2 years sexual (Spinycheek crayfish) fish opportunistic, macrophytes, benthic invertebrates Faxonius rusticus omnivore (including snails), detritus, periphyton, fish eggs, small 1–2 years sexual (Rusty crayfish) fish Filinia cornuta herbivore phytoplankton 1–7 days sexual, asexual (Wheel animal) Filinia passa herbivore bacteria 1–7 days sexual, asexual (Wheel animal) Fredericella sultana herbivore phytoplankton, bacteria 2–8 weeks sexual, asexual (Moss animal) detritus, suspended organic matter, zooplankton, Gammarus tigrinus omnivore phytoplankton, plant material, benthic 1–2 months sexual (Amphipod) macroinvertebrates Gmelinoides fasciatus detritus, sediments, phytoplankton, zooplankton, omnivore 3–4 months sexual (Baikalian amphipod) benthic macroinvertebrates Gyrodactylus salaris ectoparasite fish parasite 1 day sexual (Salmon fluke) opportunistic omnivore, zooplankton, detritus, Hemimysis anomala omnivore phytoplankton (green algae, diatoms), benthic 1–2 months sexual (Bloody red shrimp) macroinvertebrates Heterocope appendiculata omnivore zooplankton, phytoplankton 8–12 days sexual (Calanoid copepod) Heterocope caspia omnivore zooplankton, phytoplankton n/a sexual (Calanoid copepod) Hypania invalida herbivore phytoplankton (diatoms) 3 months sexual (Bristle worm) Hypanis colorata (Clam, coloured lagoon herbivore phytoplankton (green algae, diatoms), detritus 1 year sexual cockle) Lernaea cyprinacea ectoparasite fish parasite 14–90 days sexual (Anchor worm)

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Species Age to Trophic guild Diet details Reproduction (common name) maturity Leyogonimus polyoon endoparasite benthic invertebrate parasite, bird parasite n/a sexual, asexual (Trematode flatworm, leo) Limnomysis benedeni omnivore, organic matter, phytoplankton, detritus, benthic algae, (Opossum shrimp, Caspian 3–10 months sexual herbivore biofilms slender shrimp) Limnoperna fortunei omnivore phytoplankton, zooplankton 1 year sexual (Golden mussel) Melanoides tuberculatus omnivore, periphyton, detritus, diatoms, epiphytic algae, plant (Malaysian trumpet snail, red- 3–6 months sexual, asexual herbivore material, fish eggs rimmed melania) Obesogammarus crassus generalist, detritus, zooplankton, phytoplankton, omnivore n/a sexual (Scud, thick scud) benthic macroinvertebrates, plant material Obesogammarus obesus generalist, detritus, benthic macroinvertebrates, plant omnivore <1 year sexual (Scud) material Paraleptastacus spinicauda triseta omnivore bacteria 1–3 months sexual (Harpacticoid copepod) Paramysis intermedia omnivore zooplankton <1 year sexual (Opossum shrimp) Paramysis (Metamysis) ullskyi omnivore generalist, zooplankton <1 year sexual (Opossum shrimp) Paramysis lacustris omnivore phytoplankton, zooplankton <1 year sexual (Opossum shrimp) Podonevadne trigona ovum omnivore phytoplankton, zooplankton 1–4 weeks sexual, asexual (Water flea) Polypodium hydriforme carnivore fish parasite 1–3 months sexual (Sturgeon egg parasite) Pomacea canaliculata herbivore generalist, aquatic macrophytes, detritus 2–3 years sexual (Applesnail) Pontogammarus robustoides generalist, detritus, sediments, phytoplankton, benthic omnivore 4–5 weeks sexual (Scud) macroinvertebrates Potamopyrgus antipodarum generalist, detritus, epiphytic and periphytic algae, omnivore 3–9 months sexual, asexual (New Zealand mudsnail) macrophytes, sediment

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Species Age to Trophic guild Diet details Reproduction (common name) maturity Procambarus clarkii opportunistic, macrophytes, benthic invertebrates, omnivore 2–4 months sexual (Red swamp crayfish) detritus, sediment, fish Procambarus fallax f. virginalis opportunistic, macrophytes, benthic invertebrates, (Marmorkrebs, marbled omnivore 4–9 months asexual detritus, sediment, fish crayfish) Pseudodactylogyrus anguillae ectoparasite fish parasite 1–4 weeks sexual (Eel parasite) Pseudodactylogyrus bini ectoparasite fish parasite 1–4 weeks sexual (Eel gill parasite) Radix auricularia herbivore phytoplankton, aquatic plants, benthic algae, detritus <1 year sexual (European ear snail) Schyzocotyle acheilognathi endoparasite fish parasite 40–80 days sexual, asexual (Asian tapeworm) Sinanodonta woodiana herbivore phytoplankton 1–2 years sexual (Chinese pond mussel) Valvata piscinalis herbivore phytoplankton, aquatic plants, benthic algae, detritus 1 year sexual (European valve snail) Viviparus georgianus phytoplankton, aquatic plants, benthic algae, detritus, omnivore 1–3 years sexual (Banded mystery snail) sediment, fish eggs

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Table C5. Impacts of invasion (realized or potential) for current and potential invasive aquatic invertebrates in the Great Lakes region, listed alphabetically by scientific name. n/a=data not availalble. Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, Anguillicoloides crassus predation, reduced fish growth and condition in (Eel swim-bladder n/a n/a n/a indigenous species host fish, decline in indigenous fish n/a nematode) decline, disease species, possible host death vector competition, Astacus astacus vector of crayfish virus; absence of n/a n/a n/a disease vector, USFWS (European crayfish) impact research unknown Austropotamobius competition, vector of crayfish virus; absence of pallipes n/a n/a n/a disease vector, n/a impact research (White-clawed crayfish) unknown competes with indigenous competition, communities for food and resources; Bithynia tentaculata little/no indigenous species high moderate bio-fouling; ingestion of fish with USFWS (Faucet snail) evidence decline, disease trematode parasite causes waterfowl vector, nuisance mortality No species-specific information on the trophic effect of introduced B. Brachionus leydigii little/no little/no little/no not likely ledygii; however, ≈275 species of USFWS (Wheel animal) evidence evidence evidence rotifers in the GLR; impacts are unlikely competes with indigenous competition, Bythotrephes communities (zooplankton and small little/no little/no predation, longimanus high fishes) for food and resources; USFWS evidence evidence indigenous species (Spiny waterflea) changes zooplankton community decline, nuisance structure; fouls fishing lines/nets

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous Calanipeda aquaedulcis little/no communities for food and resources; high high competition n/a (Calanoid copepod) evidence changes zooplankton community structure competes with indigenous communities (zooplankton and small Cercopagis pengoi little/no little/no competition, high fishes) for food and resources; USFWS (Fish hook waterflea) evidence evidence predation indigenous zooplankton predation; fouls fishing lines/nets Chelicorophium little/no competes with indigenous curvispinum moderate moderate competition n/a evidence communities for food (Caspian mud shrimp) Cherax destructor little/no competition, competes with indigenous moderate moderate USFWS (Yabbie, yabby) evidence unknown communities for food and resources competition, vector of crayfish virus; competes Cherax tenuimanus n/a n/a n/a disease vector, with indigenous communities for food USFWS (Marron) unknown and resources thought to be relatively benign in GLR; however, limited studies Cipangopaludina inadequate inadequate little/no competition, not indicate effects on indigenous chinensis n/a research research evidence likely gastropods in experiments, competes (Chinese mystery snail) with indigenous communities for food and resources Cipangopaludina biofouling nuisance; host to human competition, japonica inadequate inadequate inadequate parasites; potentially competes with disease vector, USFWS (Japanese trapdoor, research research research indigenous communities for food and nuisance mystery snail) resources competition, habitat biofouling nuisance; competes with alteration, indigenous clams for food and Corbicula fluminea inadequate moderate moderate indigenous species resources; decline and replacement USFWS (Asian clam) research decline, disease of indigenous clams; host to human vector, nuisance parasites; alters benthic substrates

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competes with indigenous and invasive communities for food and resources; reduces prey availability Cordylophora caspia inadequate inadequate little/no competition, for indigenous fishes; restructures n/a (Freshwater hydroid) research research evidence nuisance benthic and pelagic communities; biofouling nuisance; degrades cement and mortar structured pilings Cornigerius maeoticus inadequate little/no little/no unknown lack of knowledge USFWS (Water flea) research evidence evidence competes with indigenous communities for food and resources, competition, Cyclops kolensis little/no little/no restructures and/or dominates moderate predation, disease n/a (Cyclopoid copepod) evidence evidence zooplankton communities; possible vector intermediate host for fish and human parasites not a strong competitor, but excels at Daphnia cristata inadequate little/no moderate unknown escaping predators (planktivorous n/a (Water flea) research evidence fish) competes with indigenous competition, Daphnia lumholtzi inadequate little/no inadequate zooplankton for food and resources, indigenous species USFWS (Water flea) research evidence research restructures zooplankton decline communities competes with indigenous gammarids Dikerogammarus for food and resources; vector for inadequate little/no little/no competition, haemobaphes parasites that infect invertebrate n/a research evidence evidence disease vector (Scud, demon shrimp) intestines (effect on indigenous species unknown)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment outcompetes indigenous amphipods competition, and other macroinvertebrates for food predation, and resources; reduces community Dikerogammarus little/no little/no ecosystem change, diversity; highly aggressive predation villosus high USFWS evidence evidence indigenous species on indigenous macroinvertebrates as (Killer shrimp) decline, disease well as fish eggs/juveniles; vector for vector microsporidian parasites that may affect indigenous crustaceans competition, biofouling; decreases phytoplankton predation, habitat and zooplankton biomass and alteration, outcompetes other indigenous Dreissena polymorpha little/no ecosystem change, species for food and resources; DFO, high high (Zebra mussel) evidence indigenous species increases water clarity; USFWS decline, increase biomagnification of pollutants; whole contaminant flow, lake food web changes from nuisance phytoplankton to fish competition, biofouling; decreases phytoplankton predation, habitat and zooplankton biomass and alteration, outcompetes other indigenous Dreissena rostriformis little/no ecosystem change, species for food and resources; DFO, bugensis high high evidence indigenous species increases water clarity; USFWS (Quagga mussel) decline, increase biomagnification of pollutants; whole contaminant flow, lake food web changes from nuisance phytoplankton to fish competition, competition with indigenous species Echinogammarus predation, for food and resources, declines in little/no little/no ischnus moderate indigenous species indigenous amphipod community n/a evidence evidence (Scud) decline, disease diversity, indigenous gammarid vector predation, gammarid parasite host Echinogammarus competition, unknown, competes with indigenous trichiatus n/a n/a n/a n/a unknown gammarids for food and resources (Scud)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition with indigenous species Echinogammarus competition, little/no for food and resources, declines in warpachowskyi moderate moderate indigenous species n/a evidence indigenous amphipod community (Scud) decline diversity Ectinosoma abrau inadequate little/no little/no unknown unknown n/a (Harpacticoid copepod) research evidence evidence competition, competition with indigenous species predation, habitat for food and resources, predation on Eriocheir sinensis n/a n/a n/a alteration, indigenous species, alteration of food USFWS (Chinese mitten crab) ecosystem change, webs, increases riverbank erosion nuisance with borrowing behaviour, biofouling competition, competes with indigenous species for predation, food and resources, declines in Faxonius limosus indigenous species n/a n/a n/a indigenous crayfish communities, USFWS (Spinycheek crayfish) decline, disease possible hybridization with indigenous vector, crayfish, crayfish viral host hybridization outcompetes indigenous crayfish for competition, food and resources, declines in predation, habitat indigenous crayfish communities, alteration, changes habitat structure (reduction Faxonius rusticus ecosystem change, n/a n/a n/a in macrophyte abundance), declines USFWS (Rusty crayfish) indigenous species in macroinvertebrate and sport-fish decline, disease abundance, changes to food web vector, structure, possible hybridization with hybridization indigenous crayfish no species-specific information on the Filinia cornuta little/no inadequate trophic effect; however, ≈275 species moderate not likely n/a (Wheel animal) evidence research of rotifers in the GLR, impacts are unlikely

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment unknown, no species-specific information on the trophic effect; Filinia passa inadequate little/no little/no however, ≈275 species of rotifers in not likely, unknown n/a (Wheel animal) research evidence evidence the GLR, impacts are unlikely, develops elongated spines as predator avoidance competition with indigenous species competition, habitat for food and resources, biofouling, alteration, Fredericella sultana little/no fish parasite host (causing high high indigenous species n/a (Moss animal) evidence proliferative kidney disease (PKD) in decline, disease salmonids), declines in indigenous vector, nuisance fish species outcompetes indigenous amphipods competition, for food and resources, community predation, Gammarus tigrinus inadequate little/no little/no dominance and declines in indigenous species n/a (Amphipod) research evidence evidence indigenous amphipod communities, decline, disease fish parasite host (definitive host is vector eel) competition, competition with indigenous predation, amphipods for food and resources, Gmelinoides fasciatus n/a n/a n/a indigenous species community dominance, preys on n/a (Baikalian amphipod) decline, disease indigenous amphipods, parasite and vector disease host reduced fish growth and condition in competition, host fish, decline in indigenous fish predation, species, host death (parasite burden, Gyrodactylus salaris n/a n/a n/a indigenous species secondary infections, osmoregulatory n/a (Salmon fluke) decline, disease problems), epidemic character with vector catastrophic impact on indigenous fish species

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition with indigenous species for food and resources, reduces competition, zooplankton biomass and diversity, ecosystem change, Hemimysis anomala inadequate little/no little/no ecosystem level effects (top down indigenous species DFO (Bloody red shrimp) research evidence evidence regulation of plankton), may increase decline, disease contaminant biomagnification by vector lengthening/changing food web, may increase parasitism in fishes unknown, competition with competition, Heterocope indigenous species for food and inadequate little/no little/no increase appendiculata resources, may increase contaminant n/a research evidence evidence contaminant flow, (Calanoid copepod) biomagnification by unknown lengthening/changing food web Heterocope caspia inadequate little/no little/no unknown unknown n/a (Calanoid copepod) research evidence evidence no reports of significant impacts, Hypania invalida little/no little/no little/no not likely limited influence on the overall n/a (Bristle worm) evidence evidence evidence benthic community Hypanis colorata inadequate little/no little/no (Clam, coloured lagoon unknown unknown n/a research evidence evidence cockle) competition, reduced growth and condition in host Lernaea cyprinacea indigenous species fish, host death (parasite burden, n/a n/a n/a n/a (Anchor worm) decline, disease secondary infections, respiratory vector problems) competition, reduced growth and condition in bird Leyogonimus polyoon inadequate inadequate little/no indigenous species host, bird host death and die offs, (Trematode flatworm, n/a research research evidence decline, disease able to infect American coot and likely leo) vector other gallinules

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment outcompetes indigenous macroinvertebrate communities for competition, habitat food and resources (dominant), Limnomysis benedeni little/no alteration, declines in indigenous (Opossum shrimp, moderate moderate n/a evidence indigenous species macroinvertebrate species, changes Caspian slender shrimp) decline in habitat/sediment structure through pelletization, may change food web structure biofouling; decreases phytoplankton competition, and zooplankton biomass and predation, habitat outcompetes other indigenous alteration, species for food and resources; Limnoperna fortunei ecosystem change, high high high increases water clarity; n/a (Golden mussel) indigenous species biomagnification of pollutants; whole decline, increase lake food web changes from contaminant flow, phytoplankton to fish, negatively nuisance affects borrowing invertebrates Melanoides tuberculatus competition, outcompetes indigenous gastropods (Malaysian trumpet indigenous species for food and resources, fish parasite n/a n/a n/a USFWS snail, red-rimmed decline, disease host, human parasite host, declines in melania) vector indigenous gastropod species competes with indigenous species for Obesogammarus little/no little/no competition, food and resources, able to dominate, crassus high n/a evidence evidence predation reduces abundance of zooplankton (Scud, thick scud) community competition, may outcompete indigenous species Obesogammarus predation, for food and resources, declines in little/no little/no obesus high indigenous species indigenous amphipod species, fish n/a evidence evidence (Scud) decline, disease parasite host (Acipenserideae fish vector species)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment Paraleptastacus inadequate little/no little/no spinicauda triseta unknown unknown n/a research evidence evidence (Harpacticoid copepod) unknown, may compete with competition, indigenous zooplanktivores for food Paramysis intermedia inadequate little/no little/no increase and resources, may increase n/a (Opossum shrimp) research evidence evidence contaminant flow, contaminant biomagnification by unknown lengthening food chain unknown, competes with indigenous competition, species for food and resources, can Paramysis (Metamysis) indigenous species inadequate little/no little/no dominate indigenous mysid ullskyi decline, increase n/a research evidence evidence communities, may increase (Opossum shrimp) contaminant flow, contaminant biomagnification by unknown lengthening food chain competition, competes with indigenous species for predation, food and resources, may reduce Paramysis lacustris little/no little/no ecosystem change, zooplankton communities, may moderate n/a (Opossum shrimp) evidence evidence indigenous species increase contaminant decline, increase biomagnification by lengthening food contaminant flow chain outcompetes indigenous species for Podonevadne trigona competition, little/no little/no food and resources, can dominate ovum moderate indigenous species n/a evidence evidence zooplankton community and alter (Water flea) decline food web unknown, competes with indigenous Polypodium hydriforme competition, n/a n/a n/a fish for resources, parasitizes fish n/a (Sturgeon egg parasite) predation, unknown eggs and decreases survival

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment outcompetes indigenous species for competition, habitat food and resources, heavy grazing alteration, pressure causes loss of plants and Pomacea canaliculata ecosystem change, increased turbidity, can alter n/a n/a n/a n/a (Applesnail) indigenous species ecosystem function, decline in decline, disease indigenous species, human parasite vector, nuisance host, agricultural pest to paddy crops (e.g., wild rice, watercress) outcompetes indigenous species for competition, food and resources (aggressive, predation, habitat dominant), declines in benthic Pontogammarus alteration, little/no macroinvertebrate community robustoides moderate moderate ecosystem change, n/a evidence richness, diversity and biomass, fish (Scud) indigenous species parasite host, heavy grazing pressure decline, disease may change macrophyte vector communities unknown, competes with indigenous Potamopyrgus inadequate little/no competition, species for food and resources, low antipodarum moderate n/a research evidence nuisance, unknown digestibility to predators may affect (New Zealand mudsnail) food webs, potential biofouling competition with indigenous species for food and resources (aggressive, dominant), acts as a keystone competition, species, can restructure food webs, predation, habitat decline in indigenous species, alteration, Procambarus clarkii modification of physical-chemical moderate moderate moderate ecosystem change, USFWS (Red swamp crayfish) habitat properties, heavy grazing indigenous species pressure may change macrophyte decline, disease communities, burrowing and foraging vector, nuisance activities increase turbidity and destabilize shorelines, host to vertebrate parasites

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, competition with indigenous species predation, habitat for food and resources (aggressive, Procambarus fallax f. alteration, dominant), reproduces asexually, can virginalis n/a n/a n/a ecosystem change, restructure food webs, decline in n/a (Marmorkrebs, marbled indigenous species indigenous species, heavy grazing crayfish) decline, disease pressure may change macrophyte vector communities, parasite host Pseudodactylogyrus competition, unknown, reduced fish growth and anguillae n/a n/a n/a indigenous species n/a condition in host fish (eel), host death (Eel parasite) decline, unknown competition, Pseudodactylogyrus bini unknown, reduced fish growth and n/a n/a n/a indigenous species n/a (Eel gill parasite) condition in host fish (eel), host death decline, unknown unknown, competes with indigenous competition, Radix auricularia inadequate little/no little/no species for food and resources, host disease vector, n/a (European ear snail) research evidence evidence to many parasites (including unknown swimmer's itch) competition, reduced growth and condition in host Schyzocotyle little/no little/no little/no predation, fish; increased temperature acheilognathi n/a evidence evidence evidence indigenous species dependent mortality (especially in (Asian tapeworm) decline juvenile fish); possible host death competes with indigenous communities for food and resources; Sinanodonta woodiana competition, habitat decreases indigenous unionids, n/a n/a n/a USFWS (Chinese pond mussel) alteration modifies macroinvertebrate community structure; alters benthic substrate competition, unknown, competes with indigenous Valvata piscinalis inadequate little/no little/no indigenous species species for food and resources, n/a (European valve snail) research evidence evidence decline, disease declines in indigenous gastropod vector, unknown abundance, host to many parasites

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, unknown, predation on indigenous Viviparus georgianus inadequate little/no little/no indigenous species fish eggs, declines in largemouth n/a (Banded mystery snail) research evidence evidence decline, disease bass survival, parasite host vector, unknown

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Appendix D. Current and potential invasive aquatic plants

Table D1. Current and potential invasive aquatic plant species to the Great Lakes region (GLR), listed alphabetically by scientific name. Taxonomic family and ITIS taxonomic serial number (TSN) are listed for clarity. n/a=data not available. Organism Scientific name Common name(s) ITIS TSN Basis for listing category graminoid, shrub, Arundo donax Giant reed 41450 Proposed restriction list in GLR subshrub Azolla pinnata Mosquito fern forb/herb 181820 Proposed restriction list in GLR Butomus umbellatus Flowering rush forb/herb 38886 Ontario watch list, regulated in GLR Cabomba caroliniana Fanwort forb/herb 18408 Present, Ontario watch list Callitriche stagnalis Pond water-starwort forb/herb 32062 Regulated in GLR Lesser pond sedge, European Carex acutiformis graminoid 39477 Present, Ontario watch list lake sedge Caulerpa taxifolia (aquarium/Mediterranean Killer algae macroalgae 6974 Proposed restriction list in GLR strain) Australian stonecrop, New Crassula helmsii forb/herb 894364 Regulated in GLR Zealand pygmy weed Cylindrospermopsis raciborskii Cylindro (cyanobacteria) cyanobacteria 203690 Regulated in GLR Didymo, rock snot (invasive diatom, Didymosphenia geminata 591283 Present, regulated in GLR strain) microalgae Egeria densa Brazilian waterweed forb/herb 38972 Regulated in GLR Eichhornia azurea Anchored water hyacinth forb/herb 42622 Regulated in GLR Eichhornia crassipes Water hyacinth forb/herb 42623 Ontario watch list, regulated in GLR Enteromoprha (Ulva) Sea lettuce, green alga, grass macroalgae 6535 Regulated in GLR intestinalis kelp Fallopia japonica Japanese knotweed forb/herb 565897 Regulated in GLR Glossostigma cleistanthum Mudmat forb/herb 834093 Proposed restriction list in GLR Rough mannagrass, reed Glyceria maxima graminoid 40846 Present, Ontario watch list sweetgrass Heracleum mantegazzianum Giant hogweed forb/herb 502954 Regulated in GLR

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Organism Scientific name Common name(s) ITIS TSN Basis for listing category Houttuynia cordata Chameleon forb/herb 506518 Regulated in GLR Hydrilla verticillata Hydrilla forb/herb 38974 Ontario watch list, regulated in GLR Present and regulated in GLR, Ontario Hydrocharis morsus-ranae European frogbit forb/herb 503098 watch list Hydrocotyle ranunculoides Floating marsh pennywort forb/herb 29513 Proposed restriction list in GLR Hygrophila polysperma Indian swampweed forb/herb 182342 Regulated in GLR Ipomoea aquatica Water spinach forb/herb, vine 30759 Proposed restriction list in GLR Present and regulated in GLR, Ontario Iris pseudacorus Yellow flag iris forb/herb 43194 watch list Oxygen weed, African oxygen Lagarosiphon major forb/herb 565981 Regulated in GLR weed, African elodea Landoltia punctata Dotted duckweed forb/herb 811051 Regulated in GLR Limnobium spongia Sponge plant, frog's bit forb/herb 38948 Regulated in GLR Limnophila sessiliflora Asian marshweed forb/herb 33635 Regulated in GLR Present and regulated in GLR, Ontario Lythrum salicaria Purple loosestrife forb/herb 27079 watch list Lythrum virgatum Wanded loosestrife forb/herb 27089 Proposed restriction list in GLR Water shamrock, European Marsilea quadrifolia forb/herb 17995 Regulated in GLR waterclover Melaleuca, punk tree, paperbark Melaleuca quinquenervia tree 27228 Regulated in GLR tea tree Present; USDA: invasive across North Miscanthus sacchariflorus Amur silvergrass graminoid 503854 America Chinese silvergrass, maiden Miscanthus sinensis graminoid 41874 USDA: invasive across North America grass Arrowleaf false pickerelweed, Monochoria hastata forb/herb 503870 Regulated in GLR Leaf pondweed Monochoria vaginalis Heartshape false pickerelweed forb/herb 42628 Regulated in GLR Myosotis scorpioides Aquatic forget-me-not forb/herb 31697 Proposed restriction list in GLR Parrot's feather, parrot feather Myriophyllum aquaticum forb/herb 503904 Ontario watch list, regulated in GLR watermilfoil Myriophyllum spicatum Eurasian watermilfoil forb/herb 27039 Regulated in GLR Najas marina Spiny waternymph, spiny naiad forb/herb 39001 Proposed restriction list in GLR

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Organism Scientific name Common name(s) ITIS TSN Basis for listing category Najas minor Brittle waternymph, brittle naiad forb/herb 39002 Regulated in GLR Nasturtium officinale Watercress forb/herb 23255 Regulated in GLR Nelumbo nucifera Sacred lotus forb/herb 18400 Proposed restriction list in GLR Nitellopsis obtusa Starry stonewort macroalgae n/a Regulated in GLR CA: invasive in North America, Nymphaea mexicana Banana waterlily, yellow waterlily forb/herb 18385 Minnesota watch list Nymphoides peltata Yellow floating heart forb/herb 29998 Regulated in GLR Java waterdropwort; Vietnamese Oenanthe javanica forb/herb 506796 Proposed restriction list in GLR parsley Ottelia alismoides Duck lettuce forb/herb 38976 Regulated in GLR Phantom dinoflagellate, cell from Pfiesteria spp. microalgae 553097 Regulated in GLR hell Present, Ontario watch list, regulated in Phragmites australis australis Phragmites, common reed graminoid 1053534 GLR Pistia stratiotes Water lettuce forb/herb 42542 Ontario watch list, Regulated in GLR Present and regulated in GLR, Ontario Potamogeton crispus Curly-leaf pondweed forb/herb 39007 watch list Prymnesium parvum Golden alga microalgae 2170 Regulated in GLR Ranunculus lingua Greater spearwort forb/herb n/a Regulated in GLR Sagittaria sagittifolia Arrowhead forb/herb 38931 Regulated in GLR Salvinia auriculata Giant salvinia, eared watermoss forb/herb 18013 Ontario watch list, regulated in GLR Salvinia biloba (herzogii) Giant salvinia forb/herb 566006 Regulated in GLR Salvinia molesta Giant salvinia forb/herb 181823 Ontario watch list, regulated in GLR Solanum tampicense Wetland nightshade shrub 508061 Regulated in GLR Sparganium glomeratum Bur reed forb/herb 42322 Regulated in GLR Novel cyanobacteria (order Stigonematales spp. cyanobacteria 1375 Regulated in GLR Stigonematales) Stratiotes aloides Water soldier forb/herb 505383 Ontario watch list, regulated in GLR Trapa natans Water chestnut forb/herb 27170 Ontario watch list, regulated in GLR Typha domingensis Southern cattail forb/herb 42327 Proposed restriction list in GLR Typha laxmannii Graceful cattail forb/herb n/a Proposed restriction list in GLR Typha x glauca (T. latifolia x T. Hybrid cattail, white cattail forb/herb 42328 Ontario watch list angustifolia)

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Table D2. Geographic ranges and pathways for current and potential invasive aquatic plants to the Great Lakes region (GLR), listed alphabetically by scientific name. n/a=data not available or applicable, italics indicate potential pathways for species yet to arrive in the GLR. GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range North America, commercial Africa, Arundo donax shipping, Eurasia n/a WI n/a WI, IL, OH Eurasia, aquarium/garden (Giant reed) recreational South boating, canal America, Oceania commercial Azolla pinnata Oceania, Australia, North shipping, WI n/a n/a aquarium/garden (Mosquito fern) Asia, Africa Asia, Africa America recreational boating, canal Great Lakes commercial Butomus umbellatus (all), North shipping, Eurasia n/a IL, MN, WI 1905 aquarium/garden (Flowering rush) throughout America recreational GLR boating, canal South Great Lakes North South commercial Cabomba America, (Michigan), America, America, shipping, caroliniana North America ON 1935 ON, IL, IN, Asia, aquarium/garden North recreational (Fanwort) (south and MI, NY, OH, South America boating, canal east) PA America commercial Callitriche stagnalis North shipping, (Pond water- Europe n/a IL n/a n/a America, aquarium/garden recreational starwort) Oceania boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range Great Lakes Carex acutiformis Europe, (Michigan), North (Lesser pond sedge, Eurasia, Africa, commercial recreational ON 1951 ON America, European lake Africa, Asia Southwest shipping boating, canal (Ottawa), Eurasia sedge) Asia IN, MI Oceania, commercial Oceania, North Africa, Asia, shipping, Caulerpa taxifolia Africa, Asia, America, Central WI n/a n/a aquarium/garden recreational (Killer algae) North Eurasia, America, boating, canal, America Oceania Caribbean bait release Crassula helmsii commercial (Australian North Australia, New shipping, stonecrop, New Oceania IL, MN, WI n/a n/a America, aquarium/garden Zealand recreational Zealand pygmy Europe boating, canal weed) Great Lakes Cylindrospermopsis recreational South South (Erie), ON North commercial raciborskii IL, MI, WI 1971 boating, canal, America America (Constance America shipping (Cylindro) bait release Lake), MI Didymosphenia North Western North commercial geminata America, America, IL, IN, MI, Date North shipping, canal, bait unknown (Didymo, Rock snot, Europe, Northern MN, WI unknown America recreational release invasive strain) cryptogenic Europe boating North Ohio, Illinois America, and commercial Egeria densa Brazil, Oceania, South IL, MN, Date Mississippi shipping, (Brazilian Argentina, Asia, aquarium/garden America ON unknown River recreational waterweed) Uruguay South basins, IL, boating, canal America, IN, MN, NY Europe

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range South Central commercial Eichhornia azurea America, America, North shipping, (Anchored water IL, ON n/a n/a aquarium/garden North South America recreational hyacinth) America America boating, canal North America, commercial Eichhornia crassipes South Africa, shipping, Brazil IL n/a n/a aquarium/garden (Water hyacinth) America Oceania, recreational Asia, boating, canal Europe North North North America, America, localized Enteromoprha America, Asia, Asia, Asia, Africa, populations commercial (Ulva) intestinalis Africa, Africa, recreational Europe, IL, WI 1926 in OH, MI shipping, (Sea lettuce, green Europe, South Europe, boating, canal South (Detroit aquarium/garden alga, grass kelp) America, South America, River), NY Oceania America, Oceania Oceania Great Lakes North commercial Fallopia japonica Date (all), America, shipping, (Japanese Asia East Asia MI, ON aquarium/garden unknown throughout Europe, recreational knotweed) GLR Oceania boating, canal commercial Glossostigma shipping, Australia, New North cleistanthum Oceania WI n/a n/a aquarium/garden recreational Zealand America (Mudmat) boating, canal, bait release Glyceria maxima commercial North (Rough shipping, Eurasia Europe, Asia ON 1940 ON, WI America, aquarium/garden mannagrass, reed recreational Oceania sweetgrass) boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range commercial Heracleum North IL, MI, Date shipping, mantegazzianum Asia n/a ON, WI America, aquarium/garden OH, PA unknown recreational (Giant hogweed) Europe boating, canal Asia, commercial Houttuynia cordata Oceania, shipping, Asia n/a IL n/a n/a aquarium/garden (Chameleon) North recreational America boating, canal North America, commercial NY (Finger Africa, Hydrilla verticillata IL, IN, MI, shipping, Asia India, Korea 2011 Lakes), IN, Eurasia, aquarium/garden (Hydrilla) MN, ON recreational WI South boating, canal America, Oceania Great Lakes commercial Hydrocharis morsus- IL, MI, (Erie, Eurasia, Europe, Asia, North shipping, ranae MN, ON, 1932 Ontario), aquarium/garden Africa Africa America recreational (European frogbit) WI ON, PQ, MI, boating, canal NY, VT Hydrocotyle commercial Europe, ranunculoides North Southeastern shipping, WI n/a n/a Oceania, aquarium/garden (Floating marsh America North America recreational Asia pennywort) boating, canal commercial Hygrophila North India, East shipping, polysperma Asia IL, MN, WI n/a n/a America, aquarium/garden Asia recreational (Indian swampweed) Europe boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range North Ipomoea aquatica America, commercial (Water spinach, Africa, shipping, Asia n/a WI n/a n/a aquarium/garden Swamp morning South recreational glory) America, boating, canal Oceania Great Lakes commercial IL, MI, Iris pseudacorus (all), North shipping, Eurasia n/a MN, ON, 1886 aquarium/garden (Yellow flag iris) throughout America recreational WI GLR boating, canal Lagarosiphon major commercial (Oxygen weed, Europe, Southern IL, MI, shipping, African oxygen Africa n/a n/a Africa, aquarium/garden Africa MN, WI recreational weed, African Oceania boating, canal elodea) commercial Australia, Landoltia punctata Asia, North shipping, Southeast IL n/a n/a aquarium/garden (Dotted duckweed) Oceania America recreational Asia boating, canal commercial Limnobium spongia Europe, shipping, North (Sponge plant, frog's Southern USA IL n/a n/a North aquarium/garden recreational America bit) America boating, canal, bait release commercial Limnophila India, North shipping, sessiliflora Asia Southeast IL, MN n/a n/a America, aquarium/garden recreational (Asian marshweed) Asia Asia boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range North IL, IN, MI, Great Lakes America, Eurasia, Europe, Asia, aquarium/garden, Lythrum salicaria MN, OH, (all), Eurasia, recreational Africa, North Africa, 1939 commercial (Purple loosestrife) ON, PA, throughout Oceania, boating, canal Oceania Oceania shipping WI GLR South America commercial Lythrum virgatum Europe, North shipping, Eurasia WI n/a n/a aquarium/garden (Wanded loosestrife) Western Asia America recreational boating, canal Great Lakes (Ontario), Marsilea quadrifolia commercial ON North (Water shamrock, shipping, Eurasia Europe, Asia IL 1925 (southweste America, aquarium/garden European recreational rn), NY Eurasia waterclover) boating, canal (Finger Lakes), MI North Melaleuca America, commercial quinquenervia Australia, New Asia, shipping, (Melaleuca, punk Oceania Guinea, IL, MN n/a n/a Africa, aquarium/garden recreational tree, paperbark tea Indonesia South boating, canal tree) America, Oceania localized commercial Miscanthus Date populations North shipping, sacchariflorus Asia Eastern Asia MA aquarium/garden unknown in MI, MN, America recreational (Amur silvergrass) NY, ON, WI boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range localized North Miscanthus sinensis populations America, commercial (Chinese in IL, MI, South shipping, Asia n/a n/a 1800s aquarium/garden silvergrass, maiden NY, OH, ON America, recreational grass) (southern), Oceania, boating, canal PA Europe Monochoria hastata commercial (Arrowleaf false Asia, Southeast shipping, IL, MN n/a n/a Asia aquarium/garden pickerelweed, leaf Oceania Asia, Australia recreational pondweed) boating, canal Monochoria North commercial vaginalis America shipping, Asia n/a IL, MN n/a n/a aquarium/garden (Heartshape false (Hawaii), recreational pickerelweed) Asia boating, canal Myosotis Great Lakes commercial scorpioides (all), North shipping, Eurasia Europe, Asia WI 1886 aquarium/garden (Aquatic forget-me- throughout America recreational not) GLR boating, canal Ohio, Illinois North Myriophyllum and America, commercial aquaticum IL, MI, South Amazon River Mississippi Europe, shipping, (Parrot feather, MN, ON, 2006 aquarium/garden America basin basins, IL, Asia, recreational parrot feather WI IN, MI, MN, Africa, boating, canal watermilfoil) OH, WI Oceania Myriophyllum Great Lakes North Europe, Asia, aquarium/garden, spicatum Eurasia, IL, MI, (all), America, recreational Northern 1880 commercial (Eurasian Africa MN, WI throughout Africa, boating, canal Africa shipping watermilfoil) GLR Oceania

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range North North America, Europe, East Great Lakes Najas marina America, Africa, aquarium/garden, Asia, Africa, (Ontario?), recreational (Spiny waternymph, Eurasia, MN 1864 Eurasia, canal, commercial Oceania, throughout boating spiny naiad) Africa, South shipping North America GLR Oceania America, Oceania Great Lakes (Erie, Najas minor Europe, Asia, Michigan, aquarium/garden, Eurasia, North recreational (Brittle waternymph, Northern IL, MN, WI 1932 Huron, commercial Africa America boating brittle naiad) Africa Ontario), shipping throughout GLR North America, Great Lakes commercial Europe, North Africa, Nasturtium officinale Eurasia, (all), shipping, Africa, IL 1847 Eurasia, aquarium/garden (Watercress) Africa throughout recreational Western Asia South GLR boating, canal America, Oceania commercial Iran to China, Nelumbo nucifera Asia, North shipping, Japan, WI n/a n/a aquarium/garden (Sacred lotus) Oceania America recreational Australia boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range Great Lakes (Erie, Huron, Michigan, Nitellopsis obtusa Ontario), St. North commercial recreational Eurasia n/a IL, MI, WI 1978 (Starry stonewort) Lawrence America shipping boating, canal Seaway, ON, IN, MI, MN, NY, PA, VT, WI Western Nymphaea North commercial Mexico, Canada, mexicana North America, shipping, Southeastern CA n/a n/a aquarium/garden (Banana waterlily, America Europe, recreational USA noxious Yellow waterlily) Oceania boating, canal weed list ON North commercial Nymphoides peltata Europe, (isolated in America, shipping, (Yellow floating Eurasia Japan, China, IL, MI, WI 1930 captivity), aquarium/garden Europe, recreational heart) India PQ, NY, Oceania boating, canal OH, VT, WI Oenanthe javanica commercial (Java Asia, Date North shipping, Asia, Australia WI IL, unknown aquarium/garden waterdropwort; Oceania unknown America recreational Vietnamese parsley) boating, canal commercial Ottelia alismoides Africa, Asia, Australia, North shipping, IL, MN n/a n/a aquarium/garden (Duck lettuce) Oceania Asia, Africa America recreational boating, canal

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range commercial Pfiesteria spp. shipping, (Phantom North North n/a IL n/a n/a unknown recreational dinoflagellate, cell America America boating, canal, from hell) bait release Great Lakes North Phragmites australis (all), America, commercial australis IL, MI, WI, recreational Eurasia n/a 1869 throughout Asia, shipping, (Phragmites, ON boating, canal Great Lakes Africa, aquarium/garden common reed) region Oceania North Australia and Africa, Asia, America, Pacific South localized Asia, Islands, commercial Pistia stratiotes America, populations Africa, recreational Africa, Asia, IL, ON 2000 shipping, (Water lettuce) North in ON, OH, Europe, boating, canal Mexico, aquarium/garden America, WI South Florida, West Oceania America, Indies Oceania Great Lakes (all), North Potamogeton throughout commercial Eurasia, Eurasia, North IL, MI, America, crispus Great Lakes shipping, recreational Africa, Africa, MN, ON, 1879 South (Curly-leaf region, ON aquarium/garden, boating, canal Oceania Australia WI America, pondweed) (Georgian escaped culture Oceania Bay-Severn River area) North America, unknown, Africa, recreational Prymnesium parvum unknown, cryptogenic, commercial IL, WI n/a n/a Eurasia, boating, canal, (Golden alga) cryptogenic Eastern shipping South bait release hemisphere America, Oceania

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range commercial Europe, North Ranunculus lingua shipping, Eurasia Middle East, IL n/a n/a America, aquarium/garden (Greater spearwort) recreational Russia Europe boating, canal North commercial America, Sagittaria sagittifolia shipping, Eurasia Eurasia IL, MN n/a n/a South aquarium/garden (Arrowhead) recreational America, boating, canal Oceania North South and commercial Salvinia auriculata North America, Central IL, MI, shipping, (Giant salvinia, n/a n/a America, aquarium/garden South America, MN, ON recreational eared watermoss) Asia America Mexico boating, canal commercial South and Salvinia biloba South shipping, Central IL, MI, MN n/a n/a Asia aquarium/garden (Giant salvinia) America recreational America boating, canal North America, commercial Africa, Salvinia molesta South IL, MI, shipping, Brazil n/a n/a Eurasia, aquarium/garden (Giant salvinia) America MN, ON recreational South boating, canal America, Oceania North Solanum North America, commercial tampicense America, Mexico, South Asia, shipping, IL, MN n/a n/a aquarium/garden (Wetland South America Africa, recreational nightshade) America Europe, boating, canal Oceania

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range commercial Sparganium Eurasia, Great Lakes North Europe, Asia, shipping, glomeratum North IL, MN 1936 (Superior), America, aquarium/garden North America recreational (Bur reed) America MN, WI Oceania boating, canal commercial Stigonematales spp. shipping, unknown, unknown, North (Novel IL n/a n/a unknown recreational novel novel America cyanobacteria) boating, canal, bait release ON (Trent- commercial IL, MI, North Stratiotes aloides Severn shipping, Eurasia Europe, Asia MN, ON, 2008 America, aquarium/garden (Water soldier) Waterway, recreational WI Europe Black River) boating, canal Great Lakes commercial IL, MI, North Trapa natans Eurasia, Europe, Asia, (Ontario), shipping, MN, NY, 1942 America, aquarium/garden (Water chestnut) Africa Africa PQ, NY, recreational ON, WI Oceania PA, VT boating, canal North North North America, America, localized America, Asia, Asia, commercial Asia, Africa, populations Typha domingensis Africa, indigeno Africa, shipping, Europe, WI (may be aquarium/garden (Southern cattail) Europe, South us Europe, recreational South indigenous) America, South boating, canal America, in IL, OH Oceania America, Oceania Oceania aquarium/garden, Typha laxmannii recreational Eurasia Europe, Asia WI n/a n/a Europe commercial (Graceful cattail) boating, canal shipping

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GLR Current Species GLR Current Introduction GLR spread Origin Origin detail regulated invaded (common name) year GLR range pathway pathway locations range localized populations commercial Typha x glauca North Date in IL, IN, MI, shipping, (Hybrid cattail, white Europe n/a ON America, aquarium/garden unknown MN, NY, recreational cattail) Europe OH, ON, boating, canal PQ, VT, WI

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Table D3. Size and habitat preferences of current and potential invasive aquatic plants to the Great Lakes region, listed alphabetically by scientific name. n/a=data not available. Maximum Species Salinity Habitat adult Habitat description (common name) tolerance preference length Arundo donax freshwater, 600 cm wetland, riparian wet, drained soil (Giant reed) brackish lake/pond, Azolla pinnata 2.5 cm freshwater river/stream, quiet, slow-moving water, moist soil (Mosquito fern) riparian lake/pond, Butomus umbellatus 150 cm freshwater river/stream, wet soil, water up to 2 m deep (Flowering rush) wetland, riparian Cabomba caroliniana lake/pond, 10 m freshwater still to slow-moving water (Fanwort) river/stream lake/pond, Callitriche stagnalis 30 cm freshwater river/stream, shallow, still to slow-moving water, wet soil (Pond water-starwort) wetland, riparian Carex acutiformis lake/pond, moist soil, most wetland types, margins of rivers, (Lesser pond sedge, European lake 75 cm freshwater river/stream, streams, lakes, ponds sedge) wetland, riparian 5–80 cm Caulerpa taxifolia brackish, lake/pond, epibenthic (attaches to most substrates including (primary (Killer algae) marine river/stream sand, rock, mud), shallow or deep water frond) Crassula helmsii lake/pond, wide range of habitats, terrestrial form: moist soil, 10–130 cm freshwater, (Australian stonecrop, New Zealand river/stream, most wetland types, margins of water; aquatic (stem) brackish pygmy weed) wetland, riparian form: can establish in water up to 3 m deep Cylindrospermopsis raciborskii lake/pond, planktonic: euphotic zone, prefers still to slow- 311 um freshwater (Cylindro) river/stream moving water Didymosphenia geminata lake/pond, epilithic and epiphytic, prefers cold, flowing 140 um freshwater (Didymo, rock snot, invasive strain) river/stream oligotrophic water Egeria densa lake/pond, still or flowing water up to 7 m deep, tolerates 500 cm freshwater (Brazilian waterweed) river/stream turbidity and low light

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Maximum Species Salinity Habitat adult Habitat description (common name) tolerance preference length lake/pond, wide habitat range, prefers soft substrates (mud) Eichhornia azurea 100 cm freshwater river/stream, with seasonal floods or open and slow-moving (Anchored water hyacinth) wetland, riparian water lake/pond, wide habitat range, prefers eutrophic slow flowing Eichhornia crassipes (Water hyacinth) 120 cm freshwater river/stream, water wetland, riparian freshwater, wide range of aquatic habitats, able to take Enteromoprha (Ulva) intestinalis lake/pond, 20 cm brackish, different forms - epiphytic, epilithic, floating sheets, (Sea lettuce, green alga, grass kelp) river/stream marine floating ropes, attached mats, bottle brush Fallopia japonica prefers open areas with full sun, tolerates wide 300 cm freshwater wetland, riparian (Japanese knotweed) range of soil and moisture conditions lake/pond, muddy and sandy substrates of littoral zones up to Glossostigma cleistanthum 2 cm freshwater river/stream, 4 m deep, prefers full sun and oligotrophic (Mudmat) wetland, riparian conditions Glyceria maxima lake/pond, wide range of habitats, prefers wet or occasionally (Rough mannagrass, reed 250 cm freshwater river/stream, flooded areas, slow to still water up to 30 cm deep, sweetgrass) wetland, riparian full sun to partial shade Heracleum mantegazzianum 600 cm freshwater wetland, riparian moist to wet soil, wide range of habitats (Giant hogweed) lake/pond, Houttuynia cordata terrestrial and aquatic, prefers moist loamy soils, 80 cm freshwater river/stream, (Chameleon) shallow water, shade to full sun wetland, riparian lake/pond, Hydrilla verticillata freshwater, wide range of habitats, water up to 6 m deep, 750 cm river/stream, (Hydrilla) brackish shade to full sun wetland 30 cm lake/pond, Hydrocharis morsus-ranae (individual freshwater river/stream, prefers shallow, slow-moving mesotrophic water (European frogbit) rosette) wetland lake/pond, Hydrocotyle ranunculoides wide range of habitats, prefers shallow, still to 90 cm freshwater river/stream, (Floating marsh pennywort) slow-moving water wetland

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Maximum Species Salinity Habitat adult Habitat description (common name) tolerance preference length lake/pond, Hygrophila polysperma wide range of habitats, terrestrial form can grow in 180 cm freshwater river/stream, (Indian swampweed) moist soil, aquatic form in water up to 3 m deep wetland, riparian Ipomoea aquatica lake/pond, (Water spinach, swamp morning 300 cm freshwater river/stream, wide range of habitats, moist soil or shallow water glory) wetland, riparian lake/pond, Iris pseudacorus 150 cm freshwater river/stream, wide range of habitats, moist soil or shallow water (Yellow flag iris) wetland, riparian Lagarosiphon major lake/pond, prefers slow to still water with soft substrates (Oxygen weed, African oxygen weed, 700 cm freshwater river/stream, (sand, silt), grows up to 6.6 m deep African elodea) wetland lake/pond, Landoltia punctata quiet, nutrient rich water, tolerates seasonally 10 cm freshwater river/stream, (Dotted duckweed) intermittent water wetland lake/pond, Limnobium spongia slow to still eutrophic water, wet, sandy loam soils, 2.5 cm freshwater river/stream, (Sponge plant, frog's bit) full to partial sun wetland lake/pond, wide range of habitats, wet soil to fully submersed, Limnophila sessiliflora 370 cm freshwater river/stream, cool streams to warm lake shorelines up to 3.7 m (Asian marshweed) wetland deep, full sun to shade Lythrum salicaria wide range of habitats, prefers moist rich soils, full 250 cm freshwater wetland, riparian (Purple loosestrife) sun to partial shade Lythrum virgatum wide range of habitats, prefers moist rich soils, full 100 cm freshwater wetland, riparian (Wanded loosestrife) sun to partial shade Marsilea quadrifolia lake/pond, moist soils, shallow still to slow-moving water, (Water shamrock, European 20 cm freshwater river/stream, prefers sandy and loamy soils, full sun to partial waterclover) wetland shade Melaleuca quinquenervia freshwater, (Melaleuca, punk tree, paperbark tea 12 m wetland, riparian moist soils, tolerates flooding and standing water brackish tree)

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Maximum Species Salinity Habitat adult Habitat description (common name) tolerance preference length Miscanthus sacchariflorus moist (not saturated), organic soil, full sun to partial 250 cm freshwater riparian (Amur silvergrass) shade Miscanthus sinensis moist (not saturated), organic soil, full sun to partial 370 cm freshwater riparian (Chinese silvergrass, maiden grass) shade Monochoria hastata lake/pond, (Arrowleaf false pickerelweed, leaf 100 cm freshwater river/stream, permanently wet soils, shallow still water, full sun pondweed) wetland, riparian lake/pond, Monochoria vaginalis 50 cm freshwater river/stream, permanently wet soils, shallow still water, full sun (Heartshape false pickerelweed) wetland, riparian lake/pond, Myosotis scorpioides permanently wet soils, shallow still to slow-moving 60 cm freshwater river/stream, (Aquatic forget-me-not) water wetland, riparian Myriophyllum aquaticum freshwater, lake/pond, (Parrot feather, parrot feather 150 cm wet soils, shallow water, prefers eutrophic water brackish river/stream watermilfoil) Myriophyllum spicatum freshwater, lake/pond, slow to still water up to 10 m deep, thrives in 300 cm (Eurasian watermilfoil) brackish river/stream disturbance lake/pond, Najas marina freshwater, still to slow-moving mesotrophic or eutrophic water 50 cm river/stream, (Spiny waternymph, spiny naiad) brackish up to 6 m deep wetland lake/pond, Najas minor freshwater, 120 cm river/stream, still to slow-moving water up to 4 m deep (Brittle waternymph, brittle naiad) brackish wetland lake/pond, Nasturtium officinale 60 cm freshwater river/stream, shallow, slow to fast-moving water (Watercress) wetland, riparian lake/pond, Nelumbo nucifera shallow still to slow-moving water, up to 2 m deep, 200 cm freshwater river/stream, (Sacred lotus) full sun wetland Nitellopsis obtusa freshwater, lake/pond, 200 cm deep slow-moving water, prefers soft substrates (Starry stonewort) brackish river/stream

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Maximum Species Salinity Habitat adult Habitat description (common name) tolerance preference length Nymphaea mexicana lake/pond, 150 cm freshwater deep still to slow-moving water (Banana waterlily, yellow waterlily) river/stream lake/pond, Nymphoides peltata slow-moving water 0.5-4 m deep, can survive in 200 cm freshwater river/stream, (Yellow floating heart) wet soil wetland, riparian Oenanthe javanica (Java waterdropwort; Vietnamese 100 cm freshwater wetland, riparian shallow water to moist soils, full sun to light shade parsley) lake/pond, Ottelia alismoides shallow water (5 cm to 1 m deep) to moist soils, full 70 cm freshwater river/stream, (Duck lettuce) sun to light shade wetland, riparian Pfiesteria spp. lake/pond, 60 um brackish pelagic, warm, still, eutrophic water (Phantom dinoflagellate, cell from hell) river/stream Phragmites australis australis freshwater, 600 cm wetland, riparian moist soils (Phragmites, common reed) brackish Pistia stratiotes freshwater, lake/pond, 50 cm still to slow-moving water (Water lettuce) brackish river/stream lake/pond, Potamogeton crispus freshwater, wide range of habitats and conditions (nutrients, 500 cm river/stream, (Curly-leaf pondweed) brackish water flow, temperature, light) wetland freshwater, lake/pond, Prymnesium parvum 10 um brackish, river/stream, shallow and deep water (Golden alga) marine wetland lake/pond, Ranunculus lingua prefers moist soil and shallow still to slow-moving 120 cm freshwater river/stream, (Greater spearwort) water, full sun wetland, riparian lake/pond, Sagittaria sagittifolia prefers nutrient rich muddy substrate with slow 100 cm freshwater river/stream, (Arrowhead) flowing or still water up to 45 cm deep wetland, riparian

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Maximum Species Salinity Habitat adult Habitat description (common name) tolerance preference length lake/pond, Salvinia auriculata 30 cm freshwater river/stream, prefers still to slow-moving, nutrient rich water (Giant salvinia, eared watermoss) wetland lake/pond, Salvinia biloba 30 cm freshwater river/stream, prefers still to slow-moving, nutrient rich water (Giant salvinia) wetland lake/pond, Salvinia molesta 30 cm freshwater river/stream, prefers still to slow-moving, nutrient rich water (Giant salvinia) wetland Solanum tampicense 500 cm freshwater wetland, riparian shade to full sun, tolerates temporary flooding (Wetland nightshade) lake/pond, Sparganium glomeratum muddy and peaty soils and shallow water up to 30 250 cm freshwater river/stream, (Bur reed) cm deep wetland, riparian Stigonematales spp. lake/pond, 30 um freshwater epiphytic, pelagic (Novel cyanobacteria) river/stream lake/pond, Stratiotes aloides shallow, still, nutrient rich water with mud and 60 cm freshwater river/stream, (Water soldier) organic substrates up to 6.5 m deep wetland lake/pond, Trapa natans 480 cm freshwater river/stream, still to slow-moving water up to 60 cm deep (Water chestnut) wetland lake/pond, Typha domingensis freshwater, shallow, nutrient rich water to moist soil, sun to part 400 cm river/stream, (Southern cattail) brackish shade wetland, riparian lake/pond, Typha laxmannii shallow, nutrient rich water to moist soil, sun to part 150 cm freshwater river/stream, (Graceful cattail) shade wetland, riparian lake/pond, Typha x glauca freshwater, shallow, nutrient rich water to moist soil, sun to part 300 cm river/stream, (Hybrid cattail, white cattail) brackish shade, tolerant to droughts and inundation wetland, riparian

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Table D4. Growth habit and reproductive behaviour of current and invasive aquatic plants to the Great Lakes region, listed alphabetically by scientific name. n/a=data not available. Species Growth habit Life cycle Reproduction Reproduction strategy (common name) Arundo donax riparian, sexual, perennial rhizomes, fragmentation (Giant reed) emergent asexual Azolla pinnata sexual, floating leaf annual external pollination, fragmentation (Mosquito fern) asexual Butomus umbellatus sexual, external pollination, self-pollination, emergent perennial (Flowering rush) asexual rhizomes, fragmentation Cabomba caroliniana submergent, sexual, perennial self-pollination, rhizomes, fragmentation (Fanwort) floating leaf asexual Callitriche stagnalis submergent, sexual, perennial self-pollination, fragmentation (Pond water-starwort) floating leaf asexual Carex acutiformis sexual, (Lesser pond sedge, European lake riparian perennial external pollination, rhizomes asexual sedge) Caulerpa taxifolia sexual, submergent perennial rhizomes, fragmentation (Killer algae) asexual Crassula helmsii emergent, sexual, (Australian stonecrop, New Zealand submergent, free perennial self-pollination, turions, fragmentation asexual pygmy weed) floating, riparian annual, Cylindrospermopsis raciborskii submergent, free perennial asexual fragmentation (Cylindro) floating (tropical locations) Didymosphenia geminata sexual, submergent n/a fragmentation (Didymo, rock snot, invasive strain) asexual Egeria densa sexual, submergent perennial external pollination, fragmentation (Brazilian waterweed) asexual Eichhornia azurea submergent, sexual, perennial external pollination, fragmentation (Anchored water hyacinth) emergent asexual submergent, sexual, external pollination, stolons, Eichhornia crassipes (Water hyacinth) perennial emergent asexual fragmentation

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Species Growth habit Life cycle Reproduction Reproduction strategy (common name) Enteromoprha (Ulva) intestinalis submergent, free sexual, n/a fragmentation (Sea lettuce, green alga, grass kelp) floating asexual Fallopia japonica riparian, sexual, external pollination, rhizomes, annual (Japanese knotweed) emergent asexual fragmentation Glossostigma cleistanthum submergent, annual, sexual self-pollination, rhizomes, fragmentation (Mudmat) emergent perennial Glyceria maxima riparian, sexual, external pollination, rhizomes, perennial (Rough mannagrass, reed sweetgrass) emergent asexual fragmentation Heracleum mantegazzianum riparian perennial sexual external pollination, self-pollination (Giant hogweed) Houttuynia cordata emergent, sexual, perennial rhizomes, stolons, fragmenation (Chameleon) riparian asexual Hydrilla verticillata submergent, free sexual, external pollination, tubers, turions, perennial (Hydrilla) floating asexual fragmentation Hydrocharis morsus-ranae sexual, free floating perennial external pollination, stolons, turions (European frogbit) asexual Hydrocotyle ranunculoides emergent, free sexual, external pollination, stolons, perennial (Floating marsh pennywort) floating asexual fragmentation submergent, Hygrophila polysperma sexual, emergent, free perennial stolons, fragmentation (Indian swampweed) asexual floating emergent, Ipomoea aquatica annual, sexual, external pollination, self-pollination, floating leaf, (Water spinach, swamp morning glory) perennial asexual fragmentation riparian Iris pseudacorus emergent, sexual, external pollination, stolons, rhizomes, perennial (Yellow flag iris) riparian asexual fragmentation Lagarosiphon major sexual, external pollination, rhizomes, (Oxygen weed, African oxygen weed, submergent perennial asexual fragmentation African elodea) Landoltia punctata sexual, free floating perennial external pollination, stolons (Dotted duckweed) asexual Limnobium spongia emergent, sexual, external pollination, stolons, turions, perennial (Sponge plant, frog's bit) floating leaf asexual fragmentation

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Species Growth habit Life cycle Reproduction Reproduction strategy (common name) Limnophila sessiliflora submergent, annual, sexual, self-pollination, fragmentation (Asian marshweed) emergent perennial asexual Lythrum salicaria emergent, sexual, perennial external pollination, rhizomes (Purple loosestrife) riparian asexual Lythrum virgatum emergent, sexual, perennial external pollination, rhizomes (Wanded loosestrife) riparian asexual floating leaf, Marsilea quadrifolia sexual, emergent, perennial external pollination, rhizomes (Water shamrock, European waterclover) asexual submergent Melaleuca quinquenervia emergent, perennial sexual external pollination, fragmentation (Melaleuca, punk tree, paperbark tea tree) riparian Miscanthus sacchariflorus sexual, riparian perennial external pollination, rhizomes (Amur silvergrass) asexual Miscanthus sinensis sexual, riparian perennial external pollination, rhizomes (Chinese silvergrass, maiden grass) asexual Monochoria hastata emergent, annual, sexual, external pollination, stolons, (Arrowleaf false pickerelweed, Leaf riparian perennial asexual fragmentation pondweed) Monochoria vaginalis emergent, annual, sexual, external pollination, stolons, tubers, (Heartshape false pickerelweed) riparian perennial asexual fragmentation Myosotis scorpioides emergent, sexual, external pollination, stolons, perennial (Aquatic forget-me-not) riparian asexual fragmentation Myriophyllum aquaticum submergent, perennial asexual fragmentation (Parrot feather, parrot feather watermilfoil) emergent Myriophyllum spicatum sexual, external pollination, stolons, submergent perennial (Eurasian watermilfoil) asexual fragmentation (auto) Najas marina annual, sexual, submergent external pollination, fragmentation (Spiny waternymph, spiny naiad) perennial asexual Najas minor annual, sexual, submergent external pollination, fragmentation (Brittle waternymph, brittle naiad) perennial asexual Nasturtium officinale sexual, emergent perennial external pollination, stolons (Watercress) asexual

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Species Growth habit Life cycle Reproduction Reproduction strategy (common name) Nelumbo nucifera emergent, sexual, external pollination, tubers, perennial (Sacred lotus) floating leaf asexual fragmentation Nitellopsis obtusa sexual, submergent annual external pollination, bulbs, fragmentation (Starry stonewort) asexual Nymphaea mexicana emergent, sexual, perennial external pollination, stolons (Banana waterlily, yellow waterlily) floating leaf asexual Nymphoides peltata emergent, sexual, external pollination, rhizomes, perennial (Yellow floating heart) floating leaf asexual fragmentation Oenanthe javanica sexual, external pollination, stolons, emergent perennial (Java waterdropwort; Vietnamese parsley) asexual fragmentation Ottelia alismoides submergent, annual, sexual external pollination, self-pollination (Duck lettuce) emergent flower perennial Pfiesteria spp. sexual, free floating n/a fragmentation (Phantom dinoflagellate, cell from hell) asexual Phragmites australis australis sexual, external pollination, self-pollination, riparian perennial (Phragmites, common reed) asexual stolons, rhizomes, fragmentation Pistia stratiotes sexual, external pollination, stolons, free floating perennial (Water lettuce) asexual fragmentation Potamogeton crispus sexual, external pollination, rhizomes, turions, submergent perennial (Curly-leaf pondweed) asexual fragmentation Prymnesium parvum sexual, free floating n/a fragmentation (Golden alga) asexual Ranunculus lingua sexual, emergent perennial external pollination, stolons (Greater spearwort) asexual Sagittaria sagittifolia sexual, emergent perennial external pollination, stolons, tubers (Arrowhead) asexual Salvinia auriculata free floating, annual, asexual rhizomes, fragmentation (Giant salvinia, eared watermoss) emergent perennial Salvinia biloba free floating, annual, asexual rhizomes, fragmentation (Giant salvinia) emergent perennial Salvinia molesta free floating, annual, asexual rhizomes, fragmentation (Giant salvinia) emergent perennial

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Species Growth habit Life cycle Reproduction Reproduction strategy (common name) Solanum tampicense sexual, riparian perennial external pollination, fragmentation (Wetland nightshade) asexual Sparganium glomeratum emergent, sexual, perennial external pollination, rhizomes (Bur reed) riparian asexual Stigonematales spp. free floating, sexual, n/a fragmentation (Novel cyanobacteria) epiphytic asexual emergent, Stratiotes aloides sexual, submergent, free perennial stolons, turions, fragmentation (Water soldier) asexual floating Trapa natans emergent, sexual, external pollination, self-pollination, annual (Water chestnut) floating leaf asexual fragmentation Typha domingensis sexual, external pollination, self-pollination, emergent perennial (Southern cattail) asexual rhizomes, fragmentation Typha laxmannii sexual, external pollination, self-pollination, emergent perennial (Graceful cattail) asexual rhizomes, fragmentation Typha x glauca emergent, sexual, external pollination, self-pollination, perennial (Hybrid cattail, white cattail) riparian asexual rhizomes

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Table D5. Effects of invasion (realized or potential) for current and potential invasive aquatic plants in the Great Lakes region, listed alphabetically by scientific name. n/a=data not available. Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, outcompetes indigenous communities for Arundo donax habitat alteration, resources, alters indigenous vegetative n/a n/a n/a USDA (Giant reed) ecosystem structure/habitat, fire promoter, modifies change river hydrology competition, forms dense surface mats that interfere Azolla pinnata n/a n/a n/a ecosystem with recreation; decreases oxygen and n/a (Mosquito fern) change, nuisance light levels to indigenous plants forms dense stands that displace Butomus umbellatus inadequate inadequate competition, indigenous species and interfere with moderate MISP (Flowering rush) research research nuisance recreation, decreases oxygen and light levels to indigenous plants forms dense stands that displace competition, indigenous species and interfere with Cabomba caroliniana inadequate habitat alteration, recreation, decreases oxygen and light moderate moderate MISP (Fanwort) research ecosystem levels to indigenous plants, disrupts fish change, nuisance habitat and replaces indigenous food sources Callitriche stagnalis competes with indigenous communities for n/a n/a n/a competition n/a (Pond water-starwort) resources Carex acutiformis competition, little/no little/no can form dense stands that displace (Lesser pond sedge, moderate indigenous n/a evidence evidence indigenous species European lake sedge) species decline competition, can form dense stands that displace ecosystem indigenous species (algal, seagrasses, Caulerpa taxifolia change, sessile invertebrate communities) though n/a n/a n/a n/a (Killer algae) indigenous competition for food and light; toxic to species decline, herbivores; reduces indigenous species nuisance, toxic diversity and fish habitat

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, Crassula helmsii ecosystem can form dense stands that displace (Australian stonecrop, little/no change, indigenous species, reduces biodiversity, high moderate USDA New Zealand pygmy evidence indigenous decreases water quality/flow; interferes weed) species decline, with recreation nuisance habitat alteration, able to produce toxins that can be Cylindrospermopsis little/no little/no indigenous deleterious or lethal to invertebrates, fish raciborskii moderate n/a evidence evidence species decline, and mammals; affects water quality (Cylindro) nuisance, toxic (recreation and drinking water) competition, Didymosphenia ecosystem can form dense benthic mats that displace geminata little/no change, indigenous species, reduces biodiversity, moderate moderate n/a (Didymo, rock snot, evidence indigenous alters habitat, decreases water invasive strain) species decline, quality/flow; interferes with recreation nuisance forms dense stands that displace competition, indigenous species, reduces biodiversity, Egeria densa habitat alteration, n/a n/a n/a alters habitat and water quality, provides MISP (Brazilian waterweed) ecosystem poor fish habitat, interferes with recreation, change, nuisance biofouling competition, habitat alteration, forms dense floating mats that outcompete Eichhornia azurea ecosystem indigenous species, alters ecosystem and (Anchored water n/a n/a n/a change, n/a water quality, decreases biodiversity, hyacinth) indigenous interferes with recreation, biofouling species decline, nuisance

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, habitat alteration, forms dense floating mats that outcompete ecosystem indigenous species, alters ecosystem and Eichhornia crassipes high high moderate change, water quality, alters water flow, increases n/a (Water hyacinth) indigenous evapotranspiration, decreases biodiversity, species decline, interferes with recreation, biofouling nuisance competition, forms dense mats, alters ecosystems, habitat alteration, Enteromoprha (Ulva) biofouling, produces blooms/green tides ecosystem intestinalis little/no little/no that causes cascade effects through food moderate change, n/a (Sea lettuce, green evidence evidence web, reduces biodiversity, changes water indigenous alga, grass kelp) flow and chemistry, impedes commercial species decline, and recreational activities nuisance competition, potential hybridization with indigenous habitat alteration, species, forms dense monocultures, alters ecosystem ecosystems, excludes indigenous species, Fallopia japonica change, n/a n/a n/a reduces biodiversity, alters nutrient n/a (Japanese knotweed) indigenous cycling, allelopathic (chemical suppression species decline, of competitors), strong root system can hybridization, break asphalt and concrete nuisance competition, Glossostigma unknown, forms dense mats in littoral indigenous cleistanthum n/a n/a n/a habitat, outcompetes indigenous species NYS species decline, (Mudmat) for resources, reduces biodiversity unknown competition, habitat alteration, forms dense monocultures, alters Glyceria maxima ecosystem ecosystems, excludes indigenous species, little/no little/no (Rough mannagrass, moderate change, reduces biodiversity, alters nutrient NYS evidence evidence reed sweetgrass) indigenous cycling, water flow and habitat, biofouling, species decline, decreases spawning habitat for fish nuisance

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment forms dense monocultures and competition, outcompetes indigenous species, alters habitat alteration, ecosystems, excludes indigenous species, Heracleum ecosystem reduces biodiversity, alters nutrient CFIA, mantegazzianum n/a n/a n/a change, cycling, causes shoreline erosion, NYS (Giant hogweed) indigenous decreases spawning for fish, allelopathic species decline, (chemical suppression of competitors), nuisance, toxic produces sap that is phytotoxic (human health hazard) competition, Houttuynia cordata unknown, outcompetes indigenous n/a n/a n/a ecosystem n/a (Chameleon) species, alters ecosystems change, unknown forms dense monocultures, alters competition, ecosystems, excludes indigenous species, habitat alteration, reduces biodiversity across all trophic ecosystem Hydrilla verticillata levels, alters nutrient cycling, changes high high moderate change, MISP (Hydrilla) water flow, allelopathic (chemical indigenous suppression of competitors), impedes species decline, commercial and recreational activities, nuisance biofouling competition, habitat alteration, forms dense floating mats, declines in Hydrocharis morsus- ecosystem indigenous species, reduces biodiversity, little/no ranae moderate moderate change, decreases water quality and flow, MISP evidence (European frogbit) indigenous biofouling, impedes commercial and species decline, recreational activities nuisance

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, habitat alteration, forms dense floating mats or stands, Hydrocotyle ecosystem declines in indigenous species, reduces ranunculoides n/a n/a n/a change, biodiversity, decreases water quality and n/a (Floating marsh indigenous flow, biofouling, impedes commercial and pennywort) species decline, recreational activities nuisance competition, habitat alteration, forms dense floating mats or stands, Hygrophila ecosystem declines in indigenous species, reduces polysperma moderate high moderate change, biodiversity, decreases water quality and USDA (Indian swampweed) indigenous flow, biofouling, impedes commercial and species decline, recreational activities nuisance competition, habitat alteration, forms dense mats, declines in indigenous Ipomoea aquatica ecosystem species, reduces biodiversity, decreases (Water spinach, n/a n/a n/a change, n/a water quality and flow, biofouling, impedes Swamp morning glory) indigenous commercial and recreational activities species decline, nuisance competition, forms dense stands, declines in habitat alteration, indigenous species, reduces biodiversity ecosystem Iris pseudacorus (including fish and fowl), decreases water high moderate moderate change, USDA (Yellow flag iris) quality and flow, toxic to animals (skin indigenous irritation and non-fatal poisoning, reduces species decline, available plant material to herbivores) nuisance, toxic

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, habitat alteration, forms dense stands, declines in Lagarosiphon major ecosystem indigenous species, reduces biodiversity, (Oxygen weed, n/a n/a n/a change, decreases water quality and flow, MISP African oxygen weed, indigenous biofouling, impedes commercial and African elodea) species decline, recreational activities nuisance Landoltia punctata competition, n/a n/a n/a unknown, forms dense stands n/a (Dotted duckweed) unknown Limnobium spongia competition, (Sponge plant, frog's n/a n/a n/a unknown, forms dense stands USDA unknown bit) competition, unknown, forms dense stands, declines in indigenous Limnophila sessiliflora indigenous species, reduces biodiversity, n/a n/a n/a species decline, n/a (Asian marshweed) biofouling, impedes commercial and nuisance, recreational activities unknown competition, forms dense stands, alters ecosystems, habitat alteration, declines in indigenous species ecosystem Lythrum salicaria inadequate inadequate communities and biomass, reduces high change, MISP (Purple loosestrife) research research biodiversity, alters water chemistry and indigenous flow, poor habitat for indigenous birds, species decline, insects and other species, biofouling nuisance

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, habitat alteration, ecosystem unknown, research on this plant is limited change, and is assumed similar to purple Lythrum virgatum n/a n/a n/a indigenous loosestrife, may hybridize with indigenous n/a (Wanded loosestrife) species decline, Lythrum alatum (winged loosestrife) as hybridization, well as purple loosestrife nuisance, unknown Marsilea quadrifolia inadequate little/no little/no competition, unknown, conflicting reports on invasive (Water shamrock, n/a research evidence evidence unknown behaviour, capable of dense stands European waterclover) competition, forms dense stands, displaces indigenous Melaleuca habitat alteration, plants and animals, changes water flow quinquenervia ecosystem n/a n/a n/a and soil chemistry, creates fire hazards, n/a (Melaleuca, punk tree, change, converts wetlands to uplands, decreases paperbark tea tree) indigenous biodiversity species decline forms dense stands, competes with competition, Miscanthus indigenous plants, reduces biodiversity habitat alteration, sacchariflorus n/a n/a n/a and displaces indigenous plants, n/a indigenous (Amur silvergrass) hybridizes with others in genus, creates species decline fire hazards, changes nutrient cycling forms dense stands, competes with competition, Miscanthus sinensis indigenous plants, reduces biodiversity habitat alteration, (Chinese silvergrass, n/a n/a n/a and displaces indigenous plants, USDA indigenous maiden grass) hybridizes with others in genus, creates species decline fire hazards, changes nutrient cycling Monochoria hastata unknown, forms dense stands, agricultural (Arrowleaf false competition, n/a n/a n/a weed to paddy crops (e.g., wild rice, n/a pickerelweed, Leaf unknown watercress) pondweed)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, Monochoria vaginalis forms dense stands, alters ecosystems, habitat alteration, (Heartshape false n/a n/a n/a agricultural weed to paddy crops (e.g., wild n/a ecosystem pickerelweed) rice, watercress) change competition, forms dense stands, declines in Myosotis scorpioides inadequate little/no little/no indigenous indigenous plant species, toxic to (Aquatic forget-me- n/a research evidence evidence species decline, mammals (cause poor condition, liver not) toxic disease) competition, forms dense stands and rafts, alters habitat alteration, Myriophyllum ecosystems, declines in indigenous ecosystem aquaticum species communities, reduces biodiversity, high moderate moderate change, MISP (Parrot feather, Parrot alters water chemistry and flow, biofouling, indigenous feather watermilfoil) impedes commercial and recreational species decline, activities nuisance competition, forms dense stands and rafts, alters habitat alteration, ecosystems, declines in indigenous ecosystem Myriophyllum species communities, reduces biodiversity, change, spicatum high high moderate alters water chemistry and flow, biofouling, MISP indigenous (Eurasian watermilfoil) impedes commercial and recreational species decline, activities, risk of hybridization with hybridization, indigenous milfoils nuisance competition, forms dense stands, competes with Najas marina inadequate little/no little/no indigenous indigenous species for resources, reduces (Spiny waternymph, n/a research evidence evidence species decline, biodiversity, biofouling, impedes Spiny naiad) nuisance commercial and recreational activities

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment forms dense stands that shade out littoral competition, communities, outcompetes indigenous Najas minor inadequate indigenous Najas species for resources, reduces (Brittle waternymph, moderate moderate n/a research species decline, biodiversity, biofouling, alters water flow Brittle naiad) nuisance and increases sedimentation, impedes commercial and recreational activities forms dense stands, competes with Nasturtium officinale inadequate little/no competition, moderate indigenous species for resources, alters n/a (Watercress) research evidence nuisance water flow competition, forms dense stands, competes with habitat alteration, indigenous species for resources, reduces Nelumbo nucifera indigenous n/a n/a n/a community plant and invertebrate n/a (Sacred lotus) species decline, biodiversity, biofouling, alters water flow nuisance, and increases sedimentation unknown competition, forms dense mats, declines in indigenous habitat alteration, Nitellopsis obtusa little/no species, reduces biodiversity, biofouling, moderate high indigenous MISP (Starry stonewort) evidence impedes commercial and recreational species decline, activities nuisance competition, habitat alteration, forms dense stands, declines in Nymphaea mexicana ecosystem indigenous species, reduces biodiversity, (Banana waterlily, n/a n/a n/a change, n/a impedes commercial and recreational yellow waterlily) indigenous activities species decline, nuisance

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, forms dense floating mats, declines in habitat alteration, indigenous plant communities, declines in ecosystem algal populations, food web disruption, CFIA, Nymphoides peltata little/no moderate moderate change, alters water flow and chemistry, increases USDA, (Yellow floating heart) evidence indigenous sedimentation, reduces biodiversity, MISP species decline, impedes commercial and recreational nuisance activities competition, habitat alteration, ecosystem forms dense stands, declines in Oenanthe javanica change, indigenous species, reduces biodiversity, (Java waterdropwort; n/a n/a n/a n/a indigenous impedes commercial and recreational Vietnamese parsley) species decline, activities, alters water flow nuisance, unknown competition, habitat alteration, ecosystem forms dense mats, impedes commercial Ottelia alismoides n/a n/a n/a change, and recreational activities, alters water n/a (Duck lettuce) indigenous flow and chemistry, alters ecosystem species decline, nuisance Pfiesteria spp. toxic to consumers (humans, fish, (Phantom shellfish) resulting in death and/or disease, n/a n/a n/a toxic n/a dinoflagellate, cell impedes commercial and recreational and from hell) commercial activities

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment forms dense monocultures, alters competition, ecosystems, excludes indigenous species, habitat alteration, reduces biodiversity across all trophic Phragmites australis ecosystem levels, alters nutrient cycling, changes australis change, NYS, high moderate moderate water flow, allelopathic (chemical (Phragmites, common indigenous MISP suppression of competitors), impedes reed) species decline, commercial and recreational activities, hybridization, increase fire hazards, able to hybridize nuisance with indigenous lineages competition, forms dense mats, alters ecosystems, habitat alteration, excludes indigenous species, reduces ecosystem biodiversity, alters habitat for fish and Pistia stratiotes NYS, high high n/a change, invertebrates, alters nutrient cycling, (Water lettuce) MISP indigenous changes water flow, allelopathic (chemical species decline, suppression of competitors), impedes nuisance commercial and recreational activities competition, forms dense mats, alters ecosystems, habitat alteration, excludes indigenous species, reduces ecosystem biodiversity, alters habitat for fish and Potamogeton crispus change, invertebrates, alters nutrient cycling, MISP, moderate moderate moderate (Curly-leaf pondweed) indigenous changes water flow and chemistry, able to WIDNR species decline, hybridize with indigenous pondweed, hybridization, impedes commercial and recreational and nuisance commercial activities habitat alteration, ecosystem toxic (to fish, invertebrates, planktonic Prymnesium parvum change, n/a n/a n/a algae and bacteria), alters habitat and n/a (Golden alga) indigenous ecosystems, reduces biodiversity species decline, toxic Ranunculus lingua competition, unknown, competes with indigenous n/a n/a n/a n/a (Greater spearwort) unknown species for resources, forms dense stands

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, habitat alteration, competes with indigenous species for ecosystem resources, form dense stands, alters Sagittaria sagittifolia n/a n/a n/a change, ecosystem, reduces biodiversity, changes USDA (Arrowhead) indigenous water flow and chemistry, impedes species decline, commercial and recreational activities nuisance competition, forms dense mats, alters ecosystems, habitat alteration, excludes indigenous species, reduces Salvinia auriculata ecosystem biodiversity, alters habitat for fish and (Giant salvinia, eared n/a n/a n/a change, invertebrates, alters nutrient cycling, MISP watermoss) indigenous biofouling, changes water flow and species decline, chemistry, impedes commercial and nuisance recreational activities competition, forms dense mats, alters ecosystems, habitat alteration, excludes indigenous species, reduces ecosystem biodiversity, alters habitat for fish and Salvinia biloba n/a n/a n/a change, invertebrates, alters nutrient cycling, MISP (Giant salvinia) indigenous biofouling, changes water flow and species decline, chemistry, impedes commercial and nuisance recreational activities competition, forms dense mats, alters ecosystems, habitat alteration, excludes indigenous species, reduces ecosystem biodiversity, alters habitat for fish and Salvinia molesta n/a n/a n/a change, invertebrates, alters nutrient cycling, MISP (Giant salvinia) indigenous biofouling, changes water flow and species decline, chemistry, impedes commercial and nuisance recreational activities

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, habitat alteration, forms dense stands, smothers other ecosystem vegetative growth and excludes Solanum tampicense - - - change, indigenous species, reduces biodiversity, n/a (Wetland nightshade) indigenous impedes commercial and recreational species decline, activities nuisance Sparganium inadequate little/no little/no unknown, unknown, possible hybridization risk with glomeratum n/a research evidence evidence hybridization indigenous reeds (Bur reed) indigenous toxic to consumers (birds) resulting in Stigonematales spp. - - - species decline, death, decreases biodiversity of n/a (Novel cyanobacteria) toxic indigenous green algae and diatoms competition, forms dense stands or mats, alters habitat alteration, ecosystems, excludes indigenous species, ecosystem reduces biodiversity, changes water flow Stratiotes aloides little/no USDA, moderate moderate change, and chemistry, impedes commercial and (Water soldier) evidence MISP indigenous recreational activities, suspected species decline, allelopathy (chemical suppression of nuisance competitors) competition, forms dense mats, alters ecosystems, habitat alteration, shades and excludes indigenous species, OMNRF, ecosystem reduces biodiversity, changes water flow Trapa natans CFIA, moderate high moderate change, and chemistry, impedes commercial and (Water chestnut) USDA, indigenous recreational activities, allelopathic WIDNR species decline, (chemical suppression of competitors, nuisance including phytoplankton)

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Environ- Socio- Risk Species Beneficial mental economic Impact summary Impacts and possible interactions assess- (common name) effect impact impact ment competition, forms dense stands, excludes indigenous habitat alteration, species, decreases waterfowl habitat, ecosystem alters ecosystems, alters water flow and Typha domingensis change, n/a n/a n/a increases evapotranspiration, can n/a (Southern cattail) indigenous hybridize with indigenous cattail species, species decline, impedes commercial, agricultural and hybridization, recreational activities, nuisance Typha laxmannii competition, n/a n/a n/a unknown, forms dense stands n/a (Graceful cattail) unknown competition, forms dense stands, excludes indigenous habitat alteration, species, decreases indigenous plant and Typha x glauca ecosystem invertebrate biodiversity, alters water flow (Hybrid cattail, white n/a n/a n/a change, and increases evapotranspiration, can n/a cattail) indigenous hybridize with indigenous cattail species, species decline, allelopathic (chemical suppression of hybridization competitors)

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Appendices references

Aguilera, L.A. 2015. Preventing the introduction and spread of freshwater invasive invertebrates in Ontario: Assessment of the proposed invasive species act (Bill 37). School of Environmental Studies, Queen’s University, Kingston, ON. 136 p.

Balon, E.K. 1975. Reproductive guilds of fishes: a proposal and definition. Journal of the Fisheries Research Board of Canada 32: 821–864.

Hatton, E.C., J.D. Buckley, S. Fera, S. Henry, L.M. Hunt, D.A.R. Drake and T.B. Johnson. 2018. Ecological temperature metrics for invasive fishes in Ontario and the Great Lakes region. Ontario Ministry of Natural Resources and Forestry, Science and Research Branch, Peterborough, ON. Science and Research Information Report IR-15. 27 p. + append.

Herborg, L., N.E. Mandrak, B.C. Cudmore and H.J. MacIsaac. 2007. Comparative distribution and invasion risk of snakehead (Channidae) and Asian carp (Cyprinidae) species in North America. Canadian Journal of Fisheries and Aquatic Sciences 64(12): 1723–1735.

Kolar, C. and D. Lodge. 2002. Ecological predictions and risk assessment for alien fishes in North America. Science 298: 1233–1236.

Marcogliese, D.J., A.D. Gendron, J.J.H. Forest, W. Li, K. Boyce, F. El-Shehabi, D.A.R. Drake, N.E. Mandrak, J. Sherry and J.D. McLaughlin. 2016. Range expansion and molecular confirmation of the Asian fish tapeworm in the lower Great Lakes and St. Lawrence River with notes on infections in baitfish. Journal of Great Lakes Research 42(4): 819–828.

Pagnucco, K.S., G.A. Maynard, S.A. Fera, N.D. Yan, T.F. Nalepa and A. Ricciardi. 2015. The future of species invasions in the Great Lakes-St. Lawrence River basin. Journal of Great Lakes Research 41(S1): 96–107.

Snyder, R.J., L.E. Burlakova, A.Y. Karatayev and D.B. MacNeill. 2014. Updated invasion risk assessment for Ponto-Caspian fishes to the Great Lakes. Journal of Great Lakes Research 40(2): 360–369.

Straka, M., J. Spacek and P. Paril. 2015. First record of the invasive polychaete Hypania invalida (Grube, 1960) in the Czech Republic. BioInvasions Records 4(2): 87–90.

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(.1k P.R. 19 03 18) ISBN 978-1-4868-3270-5 (print) ISBN 978-1-4868-3271-2 (pdf)