See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/267883780

Ichthyofauna of the Great Lakes Basin

Conference Paper · September 2011

CITATIONS READS 0 26

5 authors, including:

Brian M. Roth Nicholas Mandrak Michigan State University University of Toronto

33 PUBLICATIONS 389 CITATIONS 173 PUBLICATIONS 2,427 CITATIONS

SEE PROFILE SEE PROFILE

Greg G Sass Thomas Hrabik Wisconsin Department of Natural Resources University of Minnesota Duluth

95 PUBLICATIONS 796 CITATIONS 68 PUBLICATIONS 1,510 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Ecological Grass Carp Risk Assessment for the Great Lakes Basin View project

All content following this page was uploaded by Greg G Sass on 14 September 2016.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. Fishes and Decapod Crustaceans of the Great Lakes Basin

Brian M. Roth, Nicholas E. Mandrak, Th omas R. Hrabik, Greg G. Sass, and Jody Peters

The primary goal of the first edition of this chapter (Coon 1994) was to provide an overview of the Laurentian Great Lakes fish community and its origins. For this edition, we have taken a slightly diff erent approach. Although we have updated the checklist of fishes in each of the Great Lakes and their watersheds, we also include a checklist of decapod crustaceans. Our decision to include decapods derives from the lack of such a list for the Great Lakes in the literature and the importance of decapods (in particular, crayfishes) for the ecology and biodiversity of streams and lakes in the Great Lakes region (Lodge et al. 1985, 2000; Perry et al. 1997). Th is most recent checklist of fish species in the Great Lakes follows similar eff orts by Christie (1974), Bailey and Smith (1981), Underhill (1986), Coon (1994), and Cudmore-Vokey and Crossman (2000). Periodic updates are necessary to catalog changes in species composition and distribution within the Great Lakes watershed. We have made a substantial eff ort to verify presence and have included records of new introductions and their outcome. Sources include the primary literature, agency reports, the United States Geological Survey (USGS), and Department of Natural Resources agents in various states. We place particular emphasis on the coregonines of Lake Superior. Th is group of fishes is beset with controversy, including discussions of the legitimacy of their current taxonomy. We also discuss the impor- tance of coregonines to the commercial fishery of Lake Superior and their current role in the ecosystem. Th e recent origins of the Great Lakes and their biotic community define the system as dynamic. We briefly review the zoogeographic origins of the fish and decapod communities and use these origins to focus on the current communities. Lastly, we discuss future prospects for the Great Lakes fish community and how current and emerging stressors on the Great Lakes could influence the fish and decapod communities.

Definitions of the Great Lakes and Their Boundaries

Th is chapter focuses on the fishes and decapods of the five Laurentian Great Lakes (Superior, Michigan, Huron, Erie, and Ontario) and their watersheds. We follow the Great Lakes Fishery Commission (GLFC) definitions for watershed boundaries. Th e Lake Superior watershed is bounded upstream by the Lake

105 106 Brian M. Roth et al.

Nipigon drainage to the eastern end of Ogoki Lake. Lake Superior is bounded downstream by the Soo Locks on the St. Marys River. Th e Soo Locks and the Straits of Mackinac from Lake Michigan form the upstream termini of Lake Huron, whose downstream terminus is the origin of the St. Clair River. Lake Michigan’s downstream termini are the Chicago Sanitary and Ship Canal (Lake Michigan) northeast of the Des Plaines/ Kankakee River confluence near Romeoville, Illinois, andthe Mackinac Straits leading to Lake Huron. Th e Chicago Sanitary and Ship Canal (CSSC), completed in 1900, was used to export wastewater from Chicago and to breach the watershed divide between the Great Lakes and Mississippi River. Th e “boundary” between the two basins is now better defined by the system of electrical barriers at the southern end of the CSSC, rather than any geological division. Lake Erie includes the St. Clair River, Lake St. Clair, and the Detroit River. Th e division between Lake Erie and Lake Ontario is Niagara Falls. Most of the Welland Canal, which connects Lakes Erie and Ontario, is considered part of Lake Ontario. Th e downstream terminus of Lake Ontario includes the St. Lawrence River east to Cornwall, Ontario. Although Lakes Ontario and Huron are connected by the Trent-Severn Waterway, the GLFC considers the Lake Ontario watershed to begin immediately to the east of Lake Simcoe with waters to the northwest, including Lake Simcoe, belonging to the Lake Huron watershed.

Zoogeographic Origins of Great Lakes Fish and Decapod Communities

Th e fishes and decapods present in the Great Lakes and their drainages represent communities organized within the last ten thousand years or so. Th e Great Lakes themselves were formed during the retreat of the Wisconsinan glacial period ca. 14,000–9,000 years before present (BP), with the faunal communities originating from populations that survived the Wisconsinan glacial period in refugia to the northwest (Beringia), south (Mississippi), southwest (Missouri), and east (Atlantic Coast; Bailey and Smith 1981; Mandrak and Crossman 1992; Underhill 1986). In particular, much of the Great Lakes fish community is thought to have originated in the Mississippi refugium and dispersed through various outlets of the proglacial Michigan and Erie basins that overflowed into the Mississippi basin. Bailey and Smith (1981) indicated that 134 of the 174 (77 percent) fish species extant in the Great Lakes at the time of their study derived from the Mississippi refugium. Fewer fish species colonized the Great Lakes from the east through the Atlantic Coastal refugium. At least eleven fish species currently in the Great Lakes likely derived from populations that existed only in the Atlantic Coastal refugium and colonized the Great Lakes through glacial outlets in the present-day St. Lawrence, Hudson, and Susquehanna River watersheds (Bailey and Smith 1981; Mandrak and Crossman 1992; Underhill 1986). As many as twenty-six other fish species likely originated from populations in both the Atlantic Coastal and Mississippi refugia (Bailey and Smith 1981; Mandrak and Crossman 1992; Underhill 1986). Populations of several cold-water fish species are hypothesized to originate in Bering Sea drainages. Broad areas of Beringia were unglaciated and covered by fresh waters, which provided a refugium for several coldwater fish species, such as Lake Trout (common and scientific names according to Nelson et al. [2004]; scientific names are provided in table 1), Arctic Grayling, Northern Pike, and many coregonines, among others (Bailey and Smith 1981; Underhill 1986; Wilson and Hebert 1996, 1998). Dispersal occurred through a series of glacial lakes that extended from Beringia southeastward to the Great Lakes (Bailey and Smith 1981; Crossman and McAllister 1986; Underhill 1986). FISHES AND DECAPOD CRUSTACEANS 107

TABLE 1. Fish Species in the Great Lakes and Their Status

SPECIES COMMON NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Ichthyomyzon castaneus Chestnut Lamprey N N N A A Ichthyomyzon fossor Northern Brook Lamprey N N N N A Ichthyomyzon unicuspis Silver Lamprey N N N N N Lampetra appendix American Brook Lamprey N N N N N Petromyzon marinus Sea Lamprey I I I I I Acipenser fulvescens Lake Sturgeon N N N N N Polyodon spathula Paddlefi sh ep ep ep ep A Lepisosteus oculatus Spotted Gar A N A N R Lepisosteus osseus Longnose Gar N N N N N Lepisosteus platostomus Shortnose Gar A N A A A Lepisosteus platyrhincus Florida Gar A A A A IF Amia calva Bowfi n A N N N N Hiodon tergisus Mooneye A N N N N Osteoglossum bicirrhosum Arawana A IF A A A Anguilla rostrata American Eel R R R R R Alosa aestivalis Blueback Herring A A A A I Alosa chrysochloris Skipjack Herring A R A R A Alosa pseudoharengus Alewife I I I I P Alosa sapidissima American Shad A IF IF IF IF Dorosoma cepedianum Gizzard Shad A N N N N Campostoma anomalum Central Stoneroller A N N N N Campostoma oligolepsis Largescale Stoneroller A N A A A Carassius auratus Goldfi sh I I I I I Clinostomus elongatus Redside Dace I N N N N Couesius plumbeus Lake Chub N N N A N Ctenopharyngodon idella Grass Carp A I R R R Cyprinella analostanus Satinfi n Shiner A A A A N Cyprinella lutrensis Red Shiner A I A A A Cyprinella spiloptera Spotfi n Shiner A N N N N Cyprinella whipplei Steelcolor Shiner A N A A A Cyprinus carpio Common Carp I I I I I Erimystax x-punctatus Gravel Chub A A A ep A Exoglossum laurae Tonguetied Minnow A A A A N Exoglossum maxilingua Cutlip Minnow A A A A N Hybognathus hankinsoni Brassy Minnow N N N N N Hybognathus regius Eastern Silvery Minnow A A A A N Hybopsis amblops Bigeye Chub A A A N A Hypophthalmichthys molitrix Silver Carp A A A A A Hypophthalmichthys nobilis Bighead Carp A IF A IF A 108 Brian M. Roth et al.

SPECIES COMMON NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Luxilus chrysocephalus Striped Shiner A N N N N Luxilus cornutus Common Shiner N N N N N Lythrurus umbratilis Redfi n Shiner A N N N N Macrhybopsis storeriana Silver Chub A A A N A Margariscus margarita Pearl Dace N N N N N Nocomis biguttatus Hornyhead Chub N N N N N Nocomis micropogon River Chub A N N N N Notemigonus crysoleucas Golden Shiner N N N N N Notropis anogenus Pugnose Shiner A N N N N Notropis ariommus Popeye Shiner A A A ep A Notropis atherinoides Emerald Shiner N N N N N Notropis bifrenatus Bridle Shiner A A A A N Notropis blennius River Shiner A N A A A Notropis boops Bigeye Shiner A A A ep A Notropis buccatus Silverjaw Shiner A N A N A Notropis buchanani Ghost Shiner A A P P A Notropis chalybaeus Ironcolor Shiner A N A A A Notropis dorsalis Bigmouth Shiner N N A N N Notropis heterodon Blackchin Shiner N N N N N Notropis heterolepsis Blacknose Shiner N N N N N Notropis hudsonius Spottail Shiner N N N N N Notropis photogenis Silver Shiner A A A N N Notropis procne Swallowtail Shiner A A A A N Notropis rubellus Rosyface Shiner N N N N N Notropis stramineus Sand Shiner N N N N N Notropis texanus Weed Shiner A ep ep A A Notropis volucellus Mimic Shiner N N N N N Opsopoeodus emiliae Pugnose Minnow A N A N A Phenacobius mirabilis Suckermouth Minnow A A A I A Phoxinus eos Northern Redbelly Dace N N N N N Phoxinus erythrogaster Southern Redbelly Dace A N A N A Phoxinus neogaeus Finescale Dace N N N N N Pimephales notatus Bluntnose Minnow N N N N N Pimephales promelas Fathead Minnow N N N N N Pimephales vigilax Bullhead Minnow A N N R A Rhinichthys atratulus Eastern Blacknose Dace A A A A N Rhinichthys cataractae Longnose Dace N N N N N Rhinichthys obtusus Western Blacknose Dace N N N N N Scardinius erythrophthalmus Rudd A IF A I I FISHES AND DECAPOD CRUSTACEANS 109

SPECIES COMMON NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Semotilus atromaculatus Creek Chub N N N N N Semotilus corporalis Fallfi sh A A A A N Tinca tinca Tench A IF A IF IF Carpiodes carpio River Carpsucker A A A I A Carpiodes cyprinus Quillback A N N N N Catostomus catostomus Longnose Sucker N N N N N Catostomus commersonii White Sucker N N N N N Erimyzon oblongus Creek Chubsucker A N A N N Erimyzon sucetta Lake Chubsucker A N N N N Hypentelium nigricans Northern Hog Sucker A N N N N Ictiobus bubalus Smallmouth Buffalo IF R A R A Ictiobus cyprinellus Bigmouth Buffalo A I I I A Ictiobus niger Black Buffalo A I I I A Lagochila lacera Harelip Sucker A A A X A Minytrema melanops Spotted Sucker A N N N A Moxostoma anisurum Silver Redhorse N N N N N Moxostoma carinatum River Redhorse A N N N N Moxostoma duquesnei Black Redhorse A N N N N Moxostoma erythurum Golden Redhorse A N N N N Moxostoma macrolepidotum Shorthead Redhorse N N N N N Moxostoma valenciennesi Greater Redhorse A N N N N Misgurnus anguillicaudatus Oriental Weatherfi sh A I I A A Myleus pacu Pacu A A IF A IF Piaractus brachypomus Pirapatinga A IF IF IF IF Pygocentrus nattereri Red Piranha A A IF A IF Ameiurus catus White Catfi sh A A A R A Ameiurus melas Black Bullhead N N N N N Ameiurus natalis Yellow Bullhead N N N N N Ameiurus nebulosus Brown Bullhead N N N N N Ictalurus punctatus Channel Catfi sh A N N N N Noturus fl avus Stonecat N N N N N Noturus gyrinus Tadpole Madtom A N N N N Noturus insignis Margined Madtom I A I A N Noturus miurus Brindled Madtom A P A N N Noturus stigmosus Northern Madtom A A A N A Panaque nigrolineatus Royal Panaque A A A IF A Pterygoplichthys pardalis Amazon Sailfi n Catfi sh A A A IF IF Pterygoplichthys sp. Sailfi n Catfi sh A A A IF A Pylodictis olivaris Flathead Catfi sh A N R N A 110 Brian M. Roth et al.

SPECIES COMMON NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Sorubim sp. Shovelnose Catfi sh IF A A A A Esox americanus vermiculatus Grass Pickerel N N N N N Esox lucius Northern Pike N N N N N Esox masquinongy Muskellunge N N N N N Esox niger Chain Pickerel A A A IF R Dallia pectoralis Alaskan Blackfi sh A A A A IF Umbra limi Central Mudminnow N N N N N Osmerus mordax Rainbow Smelt I I I I I Coregonus artedi Cisco N N N N N Coregonus clupeaformis Lake Whitefi sh N N N N N Coregonus hoyi Bloater N N N A ep Coregonus johannae Deepwater Cisco A X X A A Coregonus kiyi Kiyi N ep ep A ep Coregonus lavaretus Powan A IF A A A Coregonus moraena German Whitefi sh A IF IF A A Coregonus nigripinnis Blackfi n Cisco N X X A A Coregonus reighardi Shortnose Cisco A X X A X Coregonus zenithicus Shortjaw Cisco N ep N ep A Oncorhynchus clarkii Cutthroat Trout A A IF A A Oncorhynchus gorbuscha Pink Salmon I I I I I Oncorhynchus kisutch Coho Salmon I I I I I Oncorhynchus masou Cherry Salmon A IF A A A Oncorhynchus mykiss Rainbow Trout I I I I I Oncorhynchus nerka Sockeye Salmon A IF IF A IF Oncorhynchus tschawytscha Chinook Salmon I I I I I Prosopium coulterii Pygmy Whitefi sh N A A A A Prosopium cylindraceum Round Whitefi sh N N N A N Salmo letnica Ohrid Trout IF A A A A Salmo salar Atlantic Salmon I I I IF ep Salmo trutta Brown Trout I I I I I Salvelinus alpinus Arctic Char A IF A A IF Salvelinus fontinalis Brook Trout N N N N N Salvelinus namaycush Lake Trout N N N N N S. fontinalis X S. namaycush Splake IF A IF A A (S. fontinalis X S. namaycush) X S. namaycush Backcross Splake A A IF IF A Salmo trutta X Salvelinus fontinalis Tiger Trout A IF A A A Thymallus arcticus Arctic Grayling ep ep ep A IF FISHES AND DECAPOD CRUSTACEANS 111

SPECIES COMMON NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Aphredoderus sayanus Pirate Perch A N N N N Percopsis omiscomaycus Trout Perch N N N N N Lota lota Burbot N N N N N Labidesthes sicculus Brook Silverside P N N N N Fundulus diaphanus Banded Killifi sh A N N N N Fundulus dispar Starhead Topminnow A N A A A Fundulus notatus Blackstripe Topminnow A N A N A Gambusia affi nis Western Mosquitofi sh A R A I IF Gambusia holbrooki Eastern Mosquitofi sh A A A IF A Apeltes quadracus Fourspine Stickleback I A A A A Culaea inconstans Brook Stickleback N N N N N Gasterosteus aculeatus Threespine Stickleback I I I I N Pungitius pungitius Ninespine Stickleback N N N N N Cottus bairdii Mottled Sculpin N N N N N Cottus cognatus Slimy Sculpin N N N N N Cottus ricei Spoonhead Sculpin N N N ep ep Myoxocephalus thompsonii Deepwater Sculpin N N N N N Morone americana White Perch I I I I I Morone chrysops White Bass N N N N N Morone mississippiensis Yellow Bass A I IF A A Ambloplites rupestris Rock Bass N N N N N Enneacanthus gloriosus Bluespotted Sunfi sh A A A A I Lepomis cyanellus Green Sunfi sh N N N N N Lepomis gibbosus Pumpkinseed N N N N N Lepomis gulosus Warmouth A N N N A Lepomis humilis Orangespotted Sunfi sh A A A I A Lepomis macrochirus Bluegill N N N N N Lepomis megalotis Longear Sunfi sh A N N N N Lepomis microlophus Redear Sunfi sh A I I I A Micropterus dolomieu Smallmouth Bass N N N N N Micropterus salmoides Largemouth Bass N N N N N Pomoxis annularis White Crappie A N N N N Pomoxis nigromaculatus Black Crappie N N N N N Ammocrypta clara Western Sand Darter A N A A A Ammocrypta pellucida Eastern Sand Darter A A ep N A Etheostoma blennioides Greenside Darter A N N N N Etheostoma caeruleum Rainbow Darter A N N N N Etheostoma chlorosomum Bluntnose Darter A ep A A A Etheostoma exile Iowa Darter N N N N N 112 Brian M. Roth et al.

SPECIES COMMON NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Etheostoma fl abellare Fantail Darter N N N N N Etheostoma microperca Least Darter N N N N N Etheostoma nigrum Johnny Darter N N N N N Etheostoma olmstedi Tessellated Darter A A A A N Etheostoma spectabile Orangethroat Darter A P A N A Etheostoma zonale Banded Darter A N A N A Gymnocephalus cernuus Ruffe I I I A A Perca fl avescens Yellow Perch N N N N N Percina caprodes Logperch N N N N N Percina copelandi Channel Darter A A N N N Percina evides Gilt Darter A A A ep A Percina maculata Blackside Darter A N N N N Percina phoxocephala Slenderhead Darter A N A A A Percina shumardi River Darter A N N N A Sander canadense Sauger N N N N ep Sander vitreus glaucus Blue Pike A A A X X Sander vitreus vitreus Walleye N N N N N Aplodinotus grunniens Freshwater Drum I N N N N Astronotus ocellatus Oscar A A A IF IF Oreochromis niloticus Nile Tilapia A A A IF A Parachromis managuensis Jaguar guapote A A A A IF Tilapia buttikoferi Zebra Tilapia A A IF A A Neogobius melanostomus Round Goby I I I I I Proterorhinus marmoratus Tubenose Goby I A I I A Channus argus Northern Snakehead A IF A A A Betta splendens Betta A A IF A A Platichthys fl esus European Flounder IF A IF IF A

A = Absent; N = Native; I = Introduced (established); R = Reportedly introduced (status uncertain); IF = Introduced and failed; P = Probably native; ep = extirpated; X = extinct

Th e colonization of many decapod crustaceans in the Great Lakes basin following the Wisconsinan glacial period is more uncertain. Th e majority of decapods in the Great Lakes basin are crayfishes, although one shrimp species (Mississippi Grass Shrimp; common and scientific names according to Williams et al. [1989] and Th oma and Jezerinac [2000]; scientific names are provided in table 2) and one non-native crab (Chinese Mitten Crab) are also present (table 2). Although the colonization of decapods into the Great Lakes is likely similar to fishes, dispersal routeshave not been entirely evaluated (Page 1985; Th oma and Jezerinac 2000). As a general rule, the number of decapod species decreases toward the north and east from the Ozark Plateau of Missouri and Arkansas and, to a lesser degree, the Eastern plateaus of Kentucky, Tennessee, Georgia, and Alabama (Crandall and Templeton 1999; Page 1985). A viable hypothesis is that FISHES AND DECAPOD CRUSTACEANS 113 most decapods in the Great Lakes basin derived from populations that existed in the Mississippi and Missouri refugia and expanded north and east at the end of the Pleistocene, similar to fishes (France 1992; Page 1985; Th oma and Jezerinac 2000). Intermittent postglacial connections, such as those between the Allegheny River and the Genessee River (now in the Lake Ontario basin), may also have aided the dispersal of decapods from the Upper Ohio River drainage (Bailey and Smith 1981; Th oma and Jezerinac 2000). Some semi-terrestrial crayfish (e.g., Devil Crayfish) may have traversed over land from disjunct drainages into those that flow into the Great Lakes (Hobbs and Jass 1988). Several crayfishes are found throughout the Great Lakes basin. Species found in the watersheds of at least three Great Lakes, including Lake Ontario, include the Common, Devil, Northern Clearwater, Virile, and Calico crayfishes. Only three species of decapods are found in every state and province surrounding the Great Lakes: the Northern Clearwater, Virile, and Rusty crayfishes. Whereas the Northern Clearwater and Virile crayfishes are considered native to most of these areas, the Rusty Crayifish is native to southeast Indiana and southwest Ohio but has spread throughout the Great Lakes region through dispersal from initial introductions related to bait-buckets and macrophyte control (Lodge et al. 1985; Magnuson et al. 1975; Th oma and Jezerinac 2000). No decapods likely originated from the Beringian refugium, given that no crayfishes are native to Alaska (Hobbs 1974) and the zoogeographical centers of cambarid crayfishes (all species found east of the Rocky Mountains) are in the Ozark and Eastern Highlands (Crandall and Templeton 1999).

TABLE 2. Decapod Crustaceans of the Great Lakes Basin

SPECIES NAME COMMON OR PROPOSED NAME SUPERIOR MICHIGAN HURON ERIE ONTARIO Cambarus bartonii Common Crayfi sh N N N N Cambarus diogenes Devil Crayfi sh N N N Cambarus polychromatus Painthanded Mudbug N Cambarus robustus Big Water Crayfi sh N N N Cambarus species A Ohio Crawfi sh N Cambarus thomai Little Brown Mudbug N Fallicambarus fodiens Digger Crayfi sh N N N Orconectes immunis Calico Crayfi sh N N N Orconectes obscurus Allegheny Crayfi sh N Orconectes propinquus Northern Clearwater Crayfi sh N N N N N Orconectes rusticus Rusty Crayfi sh I I I I I Orconectes virilis Virile Crayfi sh N N N N N Orconectes sanbornii Sanborn’s Crayfi sh N Procambarus acutus White river Crayfi sh N N Procambarus clarkii Red Swamp Crayfi sh I I Procambarus gracilis Prairie Crayfi sh N Palaemonetes kadiakensis Mississippi Grass Shrimp N N Eriochir chinensis Chinese Mitten Crab IF IF IF IF

N = Native; I = Introduced; IF = Introduced and Failed 114 Brian M. Roth et al.

Checklist of Great Lakes Fishes and Decapod Crustaceans

Fishes of the Great Lakes

Two hundred and twenty fish species are native, established, reported or failed introductions, extinct, or extirpated from the Great Lakes basin (table 1). Th e total number of extant species is 177. Of these, native species comprise 79 percent of the species count (139), with the Lake Michigan watershed containing the most native species (117) and the Lake Superior watershed containing the fewest native species (72). Th e Lake Michigan and Erie watersheds contain the largest number of established introduced species (22). Lake Ontario contains the fewest introduced species (14; table 3). Sixteen species in the Great Lakes basin have expanded their range to include at least one other lake since 1994. In particular, introduced species now found in all five Great Lake watersheds include Round Goby, White Perch, Goldfish, Common Carp, Th reespine Stickleback, and Alewife. Th e Lake Michigan watershed was colonized by Oriental Weatherfish, Yellow Bass, and Ruff e. Lake Erie was colonized by Th reespine Stickleback, River Carpsucker, Black Buff alo, and Rudd. Bigmouth Buff alo were considered native to Lake Erie by Coon (1994) but are now considered introduced (Hubbs and Lagler 2004). Two species native to other Great Lakes have expanded their range to include Lake Superior, the Freshwater Drum and Margined Madtom. Black Crappie and Yellow Bullhead, considered absent in Lake Superior by Coon (1994), are now considered native to the Lake Superior watershed (Becker 1983; Cudmore-Vokey and Crossman 2000; Mandrak 2009). Several species in Table 1 were not listed in Coon (1994) but are now considered introduced into the Great Lakes watershed: Red shiner, Blueback herring, River Carpsucker, and Smallmouth Buff alo. Th e Red Shiner is found in the Lake Michigan watershed and is thought to have been introduced near Chicago sometime in the 1950s (Hubbs and Lagler 2004). River Carpsucker was present in western Lake Erie and the Maumee River as far back as 1927 (Trautman 1981) and is now considered established in the Maumee River (Hubbs and Lagler 2004). Blueback Herring was found near Oswego, New York in October 1995 (Owens et al. 1998). Th is species is established in the upper Mohawk River, but it is unknown if natural reproduction occurs in Lake Ontario proper. Th e status of this species in Lake Ontario remains uncertain, given the difficult task of separating Blueback Herring from Alewife in large catches (M. Walsh, USGS Lake Ontario Biological Station personal communication).

TABLE 3. Lake-by-Lake Breakdowns of the Status of Ichthyofauna in the Great Lakes

STATUS SUPERIOR MICHIGAN HURON ERIE ONTARIO TOTAL Native species 72 117 102 111 106 139 Established, possibly native 0 2 1 1 1 4 Introduced, established 19 22 21 22 14 34 Introduced, reproduction unlikely 1 4 3 6 4 10 Introduced, failed 5 13 13 14 14 38 Extirpations 2 6 5 7 5 15 Extinctions 0 3 3 2 2 5 Total extant species 91 141 124 134 121 177 FISHES AND DECAPOD CRUSTACEANS 115

Smallmouth Buff alo are reported in the watersheds of Lakes Erie, Michigan, and Superior by various sources, including angling reports from the Wisconsin Department of Natural Resources (DNR; http:// dnr.wi.gov), Fuller (2009), and Becker (1983). Native to the Mississippi Basin, little information exists on the abundance or reproductive status of Smallmouth Buff alo in any Great Lake. Fuller (2009) reports this species to be native to several rivers in both the Lake Michigan and Erie basins and introduced in the Ontonagon watershed of the Lake Superior basin. Becker (1983) indicated that Smallmouth Buff alo were placed into lakes within the Ontanagon watershed in the 1930s. Recent sampling on Big Lake in the Ontanagon watershed (mentioned by Becker [1983] as a recipient of Smallmouth Buff alo) failed to yield any specimens (B.M. Roth unpublished data). Trautman (1981) detailed the introduction of Smallmouth Buff alo into Lake Erie, but the species is not listed in Hubbs et al. (2004); therefore, we consider it introduced in the Great Lakes basin. Furthermore, there are taxonomic issues with buff alo fishes in the Great Lakes basin, and this species could be misidentified. Th ere is still debate whether some species are native or introduced in the Great Lakes. Table 1 lists these species as “P” and includes Alewife in Lake Ontario, Ghost Shiner in Lakes Huron and Erie, Brook Silverside in Lake Superior, and Brindled Madtom and Orangethroat Darter in Lake Michigan. Other species considered native to other Great Lakes are reported as “R” in other lakes, including Flathead Catfish in Lake Huron, Spotted Gar in Lake Ontario, and American Eel throughout the Great Lakes; it remains unclear whether these populations represent range expansions or relict native populations. A few native species may have recolonized lakes where they were thought to be extirpated. Deepwater Sculpin, thought extirpated from Lake Ontario in the mid 1970s, now regularly appears in standardized sampling (Lantry et al. 2007; Mills et al. 2003). Th e majority of Deepwater Sculpin caught are small, although it is unclear whether this represents natural reproduction or larval drift from Lake Huron (Lantry et al. 2007; Roseman et al. 1998). Shortjaw Cisco, thought to be extirpated from Lake Huron, were rediscovered there in 2003 (N.E. Mandrak unpublished data). Eighteen species are extirpated from at least one lake. Lake Erie has lost the most species (seven), and Lake Superior has lost the fewest species (two; table 3). More than one-half of the species extirpated from the Lake Erie watershed are cyprinids, a proportion higher than any other lake. In all other lakes, corego- nines dominate the list of extinct or extirpated species. For example, five coregonines have become extinct or extirpated in Lake Michigan and four in Lake Huron. Two coregonines in the Great Lakes basin are now considered extinct: the Deepwater Cisco and Shortnose Cisco. However, there is still debate whether they represent separate species or simply a single, ecophenotypically plastic species. Th e geographic center of this debate is in Lake Superior, which contains the fewest extirpations of any Great Lake.

Lake Superior Coregonines

Th e present coregonine assemblage in Lake Superior is now composed largely of Kiyi and Cisco, although Bloater is abundant in some locations and Shortjaw Cisco is also still present. Th e coregonines that dominated the lake prior to extensive fishing, habitat degradation, and the invasion of exotic species included Shortjaw Cisco, Bloater, and Kiyi. Blackfin Cisco and Shortnose Cisco were originally described as present in Lake Superior (Koelz 1929; Scott and Crossman 1973), but those Lake Superior populations have since been synonomized with the Shortjaw Cisco (Todd and Smith 1980). Currently, the Shortjaw Cisco is present in the Superior basin in Lake Nipigon and some of its tributaries. 116 Brian M. Roth et al.

Th e first significant change to the Lake Superior coregonine assemblage included the decline of the Shortjaw Cisco between 1893 and 1910 (Lawrie and Rahrer 1973). Th e decline in Shortjaw Cisco was likely attributable to a shift in fishing for this valued species from Lake Michigan, where they had already declined, to Lake Superior, where they were abundant (Koelz 1929). Th e fishery continued to focus on the Shortjaw Cisco in Lake Superior, until they were reportedly rare by 1907–1915 (Lawrie and Rahrer 1973).

Great Lakes Deepwater Coregonines

Th e Deepwater Coregonine assemblage endemic to the Great Lakes originally contained six species. Although Koelz (1929) described more species, several were later synonomized and, despite the im- portance of the Deepwater Cisco (collectively known as “chubs”) fishery, catches were rarely identified to species (Lawrie and Rahrer 1973). Two species are now considered extinct (Deepwater Cisco and Shortnose Cisco), and the remaining four species have been extirpated from at least one Great Lake basin. All known Deepwater Cisco species declined in the Great Lakes, as a result of a combination of extensive commercial fishing, habitat degradation, and the invasion of exotic species. As a result, Great Lakes food webs have undergone substantial changes. Historically, the pelagic prey fish community was dominated by coregonines then shifted to one dominated by exotic Alewife and Rainbow Smelt . However, recently, native coregonines have increased in abundance and are the most abundant planktivore in the upper Great Lakes. Based on Ontario Ministry of Natural Resources (OMNR) fish community index netting in Lake Nipigon since 1998, Blackfin Cisco consistently represented 2–6 percent of the index catch from 1998 to 2006, although other cisco species (particularly C. artedii) declined (R. Salmon, OMNR personal communication 2007). Th e present coregonine assemblage in Lake Superior is now composed largely of Kiyi and Cisco, although Bloater is abundant in some locations and Shortjaw Cisco is also still present. A recent lake-wide, multi-agency survey of off shore areas found that Kiyi, which were relatively low in abundance historically, are now the most numerous species in Lake Superior and represent approximately 43 percent of the fish community by numerical abundance (T. Hrabik unpublished data). Although there is virtually no historic information on the population size of Kiyi (identified to species; Lawrie and Rahrer 1973; Selgeby et al. 1994), there are recent data for Lake Superior (Petzold 2002). Gillnet and trawling surveys of eastern Lake Superior in 2000–2001 indicated that Kiyi comprised anywhere between 1 and 15 percent of the chub catch, and there were an estimated 2,211 tons (271–4,452 tons; 90 percent CI) of chub in depths greater than 105 meters, the preferred depths of Kiyi. Based on these estimates, there were between 22 tons and 330 tons of Kiyi in the deepest parts of the Canadian waters of eastern Lake Superior in 2000–2001.Th e remainder of the open water community is composed of Cisco (37 percent), Bloater (11 percent), Rainbow Smelt (3 percent), and other species (7 percent; T.R. Hrabik unpublished data). However, because of their larger size, Cisco represents a larger component of biomass and contribute over 70 percent of the biomass of the community. Shortjaw Cisco abundance was high in the Koelz (1929) surveys, with an average of 121 Shortjaw Cisco per net-kilometer reported across the entire lake. In contrast, Hoff and Todd (2004) averaged 0.6 Shortjaw Cisco per net-kilometer along the south shore, Petzold (2002) averaged 1.2 Shortjaw Cisco per net-kilometer in eastern waters, Pratt and Mandrak (2007) found 5.5 Shortjaw Cisco per net-kilometer in the Rossport area, and 1.2 Shortjaw Cisco per net-kilometer were caught along the north shore in 2006 (T. Pratt unpublished data). FISHES AND DECAPOD CRUSTACEANS 117

Historically, Lakes Huron and Michigan were the only lakes with all six Deepwater Cisco species and the Cisco. Th e Deepwater Cisco was the first endemic coregonine to go extinct and was last recorded in Lake Michigan in 1951 and Lake Huron in 1952; the Shortnose Cisco was last recorded in Lake Michigan in 1982 and Lake Huron in 1985 and is also likely now extinct; and the Kiyi was last recorded in Lake Michigan in 1982 and Lake Huron in 1985 and is likely extirpated (COSEWIC 2005). In Lake Huron, Shortjaw Cisco, including the synonymised C. alpenae (Todd and Smith 1980; Todd et al. 1981), comprised approximately 25 percent of the Deepwater Cisco community in the Koelz (1929) survey but were relatively uncommon in Georgian Bay waters. Similarly, collections in American waters of Lake Huron in 1956 revealed that Shortjaw Cisco comprised 19 percent of the total Deepwater Cisco catch (USGS, Great Lakes Science Center unpublished data). Only individual specimens were taken in the 1970s, and a lone individual was taken in Lake Huron in 1982 off Au Sable Point, Michigan (Todd 1985). Shortjaw Cisco were believed to be extirpated from Lake Huron until 2002, when extensive surveys around the Bruce Peninsula, Ontario, captured a few (< 20 out of 1,943 Ciscoes examined from forty-six deepwater locations) individuals identified as Shortjaw Cisco (N.E. Mandrak unpublished data). In Lake Michigan, Shortjaw Cisco followed a similar pattern to Lake Huron. Shortjaw Cisco declined from 21 percent of the chub catch in the 1930s to 2 percent by the early 1960s before disappearing from the lake completely in the 1970s (Smith 1964; Todd 1985). Bloater are currently the most abundant Deepwater Coregonine in Lakes Huron and Michigan. In Lake Huron, the Bloater harvest plummeted in the late 1960s but increased in abundance by the 1980s, owing to strong recruitment that began in the 1970s (Mohr and Ebener 2005). By the late 1980s, Bloater were the most abundant fish in the deepwater community. In Lake Michigan, Bloater constituted more than 90 percent of the ciscoes caught in experimental gillnets in 1960–1961, persisting, whereas other cisco species collapsed because of size-selective fishing and predation by Sea Lamprey on ciscoes and their predators (Brown et al. 1985). Bloater then declined in abundance from the mid-1960s through the mid-1970s, likely as a result of the explosive increase in Alewife and the development of a targeted fishery (Brown et al. 1985). By the late 1970s, the Bloater population had increased exponentially, likely as a result of the disappearance of other ciscoes, loss of its native predator, Lake Trout, and establishment of several exotic Pacific trouts and salmons that preyed on Alewife (Brown et al. 1985). By 1989, Bloater populations were estimated to be 364 kilotons, but have declined to 23–37 kilotons by the early 2000s. Causes responsible for the decline remain speculative but include a density-dependent response or high Alewife abundance in some locations (Fleischer et al. 2005; Madenjian et al. 2008). The only ciscoes known from Lake Erie are the Cisco and Shortjaw Cisco. Only the deep eastern basin of Lake Erie is suitable for Deepwater Ciscoes. Cisco was the original dominant prey in the eastern basin and was an extremely important commercial fish in the 1920s and mid-1940s. Both the Cisco and Shortjaw Cisco have declined, however, to the point at which it was considered extirpated, although occasional catches still occur and it is now consider present in very low abundances (Markham 2009). Only a few individual Shortjaw Cisco, originally identified as C. alpenae, were ever collected from Lake Erie. Approximately forty fish were first identified out of commercial catches in the 1940s, and the last was collected in 1957 (Scott and Smith 1962). No subsequent specimens have been collected (COSEWIC 2007). Th ree of the six Deepwater Ciscoes and the Cisco were originally found in Lake Ontario. Th e Deepwater Cisco fishery (not identified to species) declined in Canadian waters from 1900 onward, increased in the 1930s, and then declined to virtually zero by the 1960s (Christie 1974). Although Koelz (1929) and Christie (1974) indicated that Blackfin Cisco were present in 1800s, Todd (1985) deemed the Lake Ontario form 118 Brian M. Roth et al. invalid. Th us, the Shortnose Cisco made up 3.4 percent of all ciscoes (n = 395) caught in experimental gill nets in 1927 (Pritchard 1931) but was not collected in a similar study in 1942 (n = 899; Stone 1947). Th e Shortnose Cisco was last recorded in 1964 and is now considered extinct. In the same studies, Kiyi made up 52.8 percent of ciscoes caught in 1927, and 0.01 percent in 1942. Only a single individual was captured in 1964, the last year that it was recorded in Lake Ontario (Wells 1969). Bloater made up 21 percent of the ciscoes caught in 1927 and 98.6 percent in 1942, but were scarce in 1969 (Wells 1969). Single specimens were caught in 1972 and 1983, the last known recorded (Mills et al. 2005). Subsequent index sampling by OMNR between 1984 and 1998 (Mills et al. 2005), commercial cisco fishing, and sampling of historic sites in western Lake Ontario in 2002 and 2009 (N.E. Mandrak unpublished data) failed to capture any Deepwater Cisco specimens.

Taxonomic Uncertainty of Other Fishes

Th e taxonomy of several fish taxa in the Great Lakes basin remains unresolved. In particular, members of the Ichthyomyzon, Ictiobus, and Esox genera contain substantial taxonomic uncertainty. Ammocoetes of the three Ichthyomyzon lamprey species found in the Great Lakes are very difficult to separate from each other, although they are readily distinguishable as adults (Scott and Crossman 1973). Neave et al. (2007) concluded that ammocoetes of the parasitic Chestnut Lamprey could only be distinguished from those of the non-parasitic Northern Brook Lamprey and parasitic Silver Lamprey in larger individuals exhibiting characteristic pigmented lateral line organs; Northern Brook and Silver Lamprey ammocoetes could never be distinguished from each other. Mandrak et al. (2004) and Docker et al. (2005) found mitochondrial DNA (mtDNA) markers for Chestnut Lamprey but did not find any reliable mtDNA or nuclear markers for Northern Brook and Silver lampreys after looking at variable regions of nuclear, mitochondrial, and microsatellite DNA. Th is suggests that these two species, thought to be a satellite species pair, may actually be ecomorphotypes of the same species. Koelz (1929) described nine cisco species in the Great Lakes, including eight species thought to be endemic and to have evolved since the end of the last ice age (ca. 10,000 years bp). As the result of synonymies (Todd and Smith 1980), the total number of cisco species in the Great Lakes was reduced to seven; however, several genetic studies have failed to find mtDNA markers that distinguish the extant cisco species (Hubert et al. 2008; Turgeon and Bernatchez 2003; Turgeon et al. 1999). Th erefore, it is possible that the diff erent species may actually be ecomorphotypes, given that whitefishes (Coregonus spp.) are known to be phenotypically plastic (Lindsey 1981). Buff alo fishes Ictiobus( spp.) with two types of mouth shapes have been collected in the Great Lakes basin. Th e Bigmouth Buff alo is readily identified by its terminal mouth, but buff alo fishes with subterminal mouths may be Black Buff alo, Smallmouth Buff alo, or some admixture and cannot be readily distinguished in the Great Lakes basin. A preliminary genetic examination of buff alo specimens collected in the Great Lakes basin indicated that all individuals exhibited introgressive hybridization among all three Buff alo species (H.L. Bart, Tulane University personal communication). Th erefore, the taxonomic identity of all Buff alo fishes in the Great Lakes basin is in question. A recent barcoding study of Canadian freshwater fishes identified all of these taxonomic uncertainties, as well as several others (Hubert et al. 2008). Several species in the Great Lakes basin have the potential to contain cryptic species, including Brook Stickleback, Mottled Sculpin, and Redfin Pickerel. Redfin Pickerel FISHES AND DECAPOD CRUSTACEANS 119

(Esox americanus) has previously been found polyphyletic, based on cyt b, with one subspecies, Redfin Pickerel (E. a. americanus), monophyletic with Chain Pickerel (E. niger) and the other subspecies, Grass Pickerel (Esox a. vermiculatus), the sister group to E. a. americanus + E. niger (Grande et al. 2004). Th e results of the barcoding study may lead to the Grass Pickerel being elevated to full species status. Brook Stickleback and Mottled Sculpin may also be divided into several species based on barcoding results. Th e diff erence in species nomenclature between Hubbs et al. (2004) and Nelson et al. (2004) may cause confusion regarding the number and composition of fish species existing in the Great Lakes. Hubbs et al. (2004) elevated two subspecies of Creek Chubsucker (Erimyzon oblongus) to full species status (Western Creek Chubsucker E. claviformis and Eastern Creek Chubsucker E. oblongus) and two subspecies of Pearl Dace (Margariscus margarita) to full species status (Allegheny Pearl Dace M. margarita and Northern Pearl Dace M. nachtriebi). Hubbs et al. (2004) also elevated Blue Pike (Sander glaucus) from subspecies to full species status, recognized Ives Lake Cisco (Coregonus hubbsi) as a new species, and resurrected the Nipigon Tullibee (C. nipigon) as a species, despite being previously synonomized with Shortjaw Cisco (C. zenithicus). Furthermore, they divided sixteen species into thirty-two subspecies (table 4) and identified nineteen species to the subspecific level (Hubbs and Lagler 2004). eTh American Fisheries Society (AFS) Names Committee may not accept these species level changes in the next version of the common and scientific names list (due in 2012) without sufficient justification. At the subspecific level, the AFS Names Committee does not provide common and scientific names, the distribution of most subspecies is unknown particularly in introgression zones (e.g., Lake Ontario), and there is a lack of comprehensive keys containing subspecies in the Great Lakes basin. Th erefore, we recommend following the species-level taxonomy outlined in existing AFS Names Committee list (Nelson et al. 2004).

Identification Problems

Several keys exist for identifying fishes in the GreatLakes basin (e.g., Bailey and Smith 1981; Scott and Crossman 1973; Trautman 1981). However, in addition to those species with taxonomic uncertainties previously described, there are species in the Great Lakes basin that have resolved taxonomies but are particularly difficult to identify. Several of these species were identified in polyphyletic clades in a recent barcoding study of Canadian freshwater fishes (Hubert et al. 2008). Th e Pugnose Minnow and Pugnose Shiner, although in diff erent genera, are morphologically very similar. Th e key distinguishing character is a diff erence in anal ray count by one ray (Bailey et al. 2004), a difficult character to assess in such a small species (typically < 50 mm). Th e Ghost Shiner is very difficult to distinguish from the Mimic Shiner and typi- cally requires examination of infraorbital canals to diff erentiate between the two species (Bailey et al. 2004). Increased attention to the identification of Mimic Shiner has led to a substantial increase in the known distribution of Ghost Shiner in the Canadian Great Lakes basin (N.E. Mandrak unpublished data). As a result of this increased knowledge and a morphological study on the geographic variation in Ghost Shiner (Kott and Fitzgerald 2000), this species is likely native to the Great Lakes basin, rather than introduced as previously thought. In the past, the Striped Shiner was not recognized as a species separate from Common Shiner in the Great Lakes basin (Scott and Crossman 1973). Th is species is now recognized as present, but the two species are often difficult to distinguish because of overlapping pre-dorsal scale counts (Kott and Fitzgerald 2000) and ambiguous dorsal striping. Hubert et al. (2008) confirmed that the morphologically similar Johnny Darter and Tessellated Darter were valid species and exhibit introgression in the Lake 120 Brian M. Roth et al.

TABLE 4. Species in the Great Lakes Identifi ed to Subspecies in Hubbs et al. (2004)

NELSON ET AL. 2004 HUBBS ET AL. 2004

COMMON NAME SCIENTIFIC NAME COMMON NAME SCIENTIFIC NAME Quillback Carpiodes cyprinus Northern Quillback Carpiodes cyprinus cyprinus Central Quillback Carpiodes cyprinus hinei River Carpsucker Carpiodes carpio Northern River Carpsucker Carpiodes carpio carpio White Sucker Catostomus commersonii White Sucker Catostomus commersonii commersonii Dwarf White Sucker Catostomus commersonii utawana Longnose Sucker Catostomus catostomus Longnose Sucker Catostomus catostomus catostomus Dwarf Longnose Sucker Catostomus catostomus nannomyzon Lake Chubsucker Erimyzon sucetta Western Lake Chubsucker Erimyzon sucetta kennerlii Shorthead Redhorse Moxostoma macrolepidotum Northern Shorthead Redhorse Moxostoma macrolepidotum macrolepidotum Lake Chub Couesius plumbeus Northern Lake Chub Couesius plumbeus plumbeus Prairie Lake Chub Couesius plumbeus dissimilis Central Stoneroller Campostoma anomalum Ohio Stoneroller Campostoma anomalum anomalum Central Stoneroller Campostoma anomalum pullum Gravel Chub Erimystax x-punctatus Eastern Gravel Chub Erimystax x-punctatus trautmani Tonguetied Minnow Exoglossum laurae Eastern Tonguetied Minnow Exoglossum laurae laurae Redfi n Shiner Lythrurus umbratilis Northern Redfi n Shiner Lythrurus umbratilis cyanocephalus Common Shiner Luxilis cornutus Northern Common Shiner Luxilis cornutus frontinalis Golden Shiner Notemigonus crysoleucas Western Golden Shiner Notemigonus crysoleucas auratus Eastern Golden Shiner Notemigonus crysoleucas crysoleucas Bigmouth Shiner Notropis dorsalis Central Bigmouth Shiner Notropis dorsalis dorsalis Eastern Bigmouth Shiner Notropis dorsalis keimi Sand Shiner Notropis stramineus Northeastern Sand Shiner Notropis stramineus stramineus Fathead Minnow Pimephales promelas Fathead Minnow Pimephales promelas promelas Longnose Dace Rhinichthys cataractae Great Lakes Longnose Dace Rhinichthys cataractae cataractae Creek Chub Semotilus atromaculatus NorthernCreek Chub Semotilus atromaculatus atromaculatus Cisco Coregonus artedi Cisco Coregonus artedi artedi Kiyi Coregonus kiyi Kiyi Coregonus kiyi kiyi Ontario Kiyi Coregonus kiyi orientalis Blackfi n Cisco Coregonus nigripinnis Michigan Blackfi n Cisco Coregonus nigripinnis nigripinnis Nipigon Blackfi n Cisco Coregonus nigripinnis regalis Shortjaw Cisco Coregonus zenithicus Shortjaw Cisco Coregonus zenithicus Sisiwit Lake Cisco Coregonus zenithicus bartletti Lake Trout Salvelinus namaycush Lake Trout Salvelinus namaycush namaycush Siscowet Salvelinus namaycush siscowet Arctic Grayling Thymallus arcticus Arctic Grayling Thymallus arcticus signifer Muskellunge Esox masquinongy Great Lakes Muskellunge Esox masquinongy masquinongy Pirate Perch Aphredoderus sayanus Western Pirate Perch Aphredoderus sayanus gibbosus Banded Killifi sh Fundulus diaphanus Eastern Banded Killifi sh Fundulus diaphanus diaphanus FISHES AND DECAPOD CRUSTACEANS 121

NELSON ET AL. 2004 HUBBS ET AL. 2004

COMMON NAME SCIENTIFIC NAME COMMON NAME SCIENTIFIC NAME Western Banded Killifi sh Fundulus diaphanus menona Greenside Darter Etheostoma blennioides Greenside Darter Etheostoma blennioides blennioides Fantail Darter Etheostoma fl abellare Barred Fantail Darter Etheostoma fl abellare fl abellare Striped Fantail Darter Etheostoma fl abellare lineolatum Johnny Darter Etheostoma nigrum Central Johnny Darter Etheostoma nigrum nigrum Scaly Johnny Darter Etheostoma nigrum eulepis Orangethroat Darter Etheostoma spectabile Northern Orangethroat Darter Etheostoma spectabile spectabile Logperch Percina caprodes Ohio Logperch Percina caprodes caprodes Northern Logperch Percina caprodes semifasciata Bluegill Lepomis macrochirus Bluegill Lepomis macrochirus macrochirus Longear Sunfi sh Lepomis megalotis Northern Longear Sunfi sh Lepomis peltastes Mottled Sculpin Cottus bairdii Northern Mottled Sculpin Cottus bairdii bairdii Great Lakes Mottled Sculpin Cottus bairdii kumlieni Slimy Sculpin Cottus cognatus Slimy Sculpin Cottus cognatus gracilis

Ontario drainage, as Chapleau and Pageau (1982) previously concluded based on morphological data. However, these species and their hybrids are very difficult to distinguish in the Lake Ontario basin, as the best characters are overlapping counts of submandibular pores and soft dorsal fin rays (Chapleau and Pageau 1982). Nelson et al. (2004) elevated two subspecies of Blacknose Dace to full species status: Western Blacknose Dace (R. obtusus), found throughout the Great Lakes basin, and Eastern Blacknose Dace (R. atratulus), likely limited to the Lake Ontario basin in the Great Lakes where it may share an introgression zone with the Western Blacknose Dace (Fraser et al. 2005). Th is taxonomic elevation is supported by genetic data (Hubert et al. 2008), but Fraser et al. (2005) found the two species morphologically indistinguishable in a recent study of twenty Canadian populations, including ten populations in the Great Lakes basin.

Decapod Crustaceans of the Great Lakes

We documented a total of eighteen decapod crustacean species either native or introduced into the Great Lakes basin (table 2). Lake Erie contains the largest number of decapod species (seventeen), whereas Lake Superior contained the fewest (six). Given the lack of recent information on decapod distributions in the basin outside the watersheds of western Lake Michigan and southern Lake Erie, we were unable to determine if any decapod species have been extirpated. For example, the last systematic survey of decapods in the State of Michigan was in the late 1920s (Creaser 1931), and the most recent decapod survey in Illinois was published nearly twenty-five years ago (Page 1985). At least one crayfish species (Painthanded Mudbug, C. polychromatus) present in the Great Lakes basin has been named since these surveys. Red Swamp Crayfish has likely expanded its distribution in the Great Lakes dramatically, given its discovery in several drowned river mouths of Lake Michigan (Simon and Th oma 2006). Systematic surveys 122 Brian M. Roth et al. of decapod distributions are needed in many locales to comprehensively assess priorities for conservation or management of native and invasive decapod species.

Post-Colonial Influences on Great Lakes Fish Community

Fishing

Since the arrival of humans along the shores of the Great Lakes thousands of years ago, the fish and inver- tebrate communities of the Great Lakes have been subjected to harvest. Th e archaelogical record suggests that humans were regularly harvesting fishes from the upper Great Lakes by 5000 years BP (Cleland 1982). First Nations and Native Americans utilized hook-and-line, spears, harpoons, and gillnets to capture Walleye, Lake Whitefish, American Eel, suckers (Catostomidae), and Lake Sturgeon, among others, for subsistence. According to notes from early explorers and ethnographers, fishes were still abundant by European contact and settlement (Cleland 1982). Th e onset of commercial fishing began with the growthof cities surrounding the Great Lakes. Fishing intensified as the population of these cities grew. Lake Sturgeon all but disappeared from all five lakes by the turn of the twentieth century, as did Deepwater Cisco in Lake Superior (Christie 1974). Following the incorporation of nylon monofilament into gillnets in the mid-1900s, fisheries for Lake Trout, Walleye, Lake Whitefish, and Cisco declined precipitously in manylakes (Christie 1974). Lake Trout were already in decline from the rapid invasion of sea lamprey, and catches of this species declined to commercial extinc- tion in all five lakes by 1960 although the species never disappeared entirely (Christie 1974). In contrast, Blue Pike, which was endemic to Lakes Erie and Ontario, became extinct in the early 1980s (Underhill 1986). Th e last confirmed occurrence of this fish was from 1983 although catches approached zero by 1960. Overfishing is thought to be a major cause of their decline, although the introduction of Rainbow Smelt is also implicated (Christie 1974). In Lake Superior, the coregonine fishery shifted from a focus on Deepwater Cisco to the Shortjaw Cisco and Cisco between 1915 and 1930, owing to demand resulting from declines in the fisheries on coregonines in the lower Great Lakes (Lawrie and Rahrer 1973). By the 1930s, the Shortjaw Cisco was reportedly the only large coregonine abundant enough to support a commercial fishery (Van Oosten 1937), although Cisco was apparently still abundant. By the 1960s, the Shortjaw Cisco populations in Lake Superior were in significant decline and by the 1970s, the commercial catch was composed primarily of Bloater and Cisco (Lawrie and Rahrer 1973). A decline in the Lake Trout fishery, as a consequence of Sea Lamprey invasion, focused fishing pressure on the remaining Bloater populations in the 1970s. Bloater apparently increased as a result of lower predation pressure resulting from the decline in their dominant predator, the Lake Trout. Th e coregonine fishery focused on Bloater and Cisco populations through the 1970s. Cisco is currently the focus of summer filet and fall roe fisheries in Minnesota waters and roe fisheries in Wisconsin, Michigan, and Canadian waters.

Habitat Degradation

Habitat modification and degradation are important agents of change in Great Lakes fish communities (Kelso et al. 1996). However, assessing the extent of habitat alteration and degradation and its eff ect on FISHES AND DECAPOD CRUSTACEANS 123 fish communities in the Great Lakes basin remains a daunting task. Habitat assessments can include physical (e.g., temperature, rocky reefs), biological (e.g., macrophytes), and chemical (e.g., nutrients) elements, all of which are dynamic in nature and potentially linked. Th ese linkages are critical in wetlands, which are potentially influenced by their adjacent terrestrial and aquatic ecosystems. Wetlands around the Great Lakes have been greatly aff ected by anthropogenic activities, such as damming, dredging, road building, draining, and nutrient loading, among others, that can lead to wetland loss and degradation (Brazner 1997; Dodge and Kavetsky 1994). Estimates of wetland loss vary by location but can be as high as 90 percent in Wetlands and nearshore habitats provide important habitat for many Great Lake fishes, owing primarily to the abundance of aquatic macrophytes in these areas (Brazner 1997; Randall et al. 1996). In turn, macrophyte cover is positively correlated with fish abundance and biomass in the Great Lakes (Randall et al. 1996). Wetlands support high fish diversity and are important as nursery areas for some species (Brazner 1997; Dodge and Kavetsky 1994; Randall et al. 1996). More pristine nearshore habitats support higher genetic diversity of brown bullhead than impaired areas (Murdoch and Hebert 1994) and fewer non-native species than impaired and heavily modified areas (Bhagat et al. 2007; Minns et al. 1994). Fish that inhabit open water areas have experienced less direct manipulation of habitat. However, chemical and biological stressors remain important influences on open water habitat quality. In western and central Lake Erie, eutrophication has led to a hypoxic “Dead Zone” that has persisted almost annually for more than fifty years during late summer (Carrick et al. 2005; Charlton and Milne 2004; Edwards et al. 2005). Declines in Hexagenia mayfly nymphs and other large benthic invertebrates linked to the hypoxia likely played a role in the decline in Yellow Perch abundance and growth from the 1960s to the early 1980s (Bridgeman et al. 2006; Tyson and Knight 2001). Although the introduction and subsequent proliferation of zebra mussels have lessened the appearance of cultural eutrophication in Lake Erie and its eff ect on Hexagenia, Yellow Perch recruitment remains sporadic (Tyson and Knight 2001). Eutrophication was instigated by the use of phosphate-based laundry detergent and bolstered by agricultural runoff . Algal blooms are frequently observed in Green Bay on Lake Michigan, Saginaw Bay on Lake Huron, and the Bay of Quinte on Lake Ontario. Eutrophication can lead to a change in aquatic communities, productivity, trophic structure, and habitat, all of which are apparent at various locations in and around the Great Lakes. Eutrophication may also negatively aff ect the quality and quantity of off shore spawning reefs for several species, including Lake Trout. Siltation derived from eutrophication may reduce the suitability of substrate for Lake Trout spawning (Sly and Evans 1996). Nonetheless, much of the physical structures (e.g., bedrock outcroppings) that support Lake Trout spawning throughout the Great Lakes are still viable sites for egg deposition (Edsall and Kennedy 1995). Non-native interstitial predators now abundant on this substrate may prohibit successful reproduction of Lake Trout in some areas of the Great Lakes by preying on Lake Trout eggs, particularly in Lake Michigan (Claramunt et al. 2005; Jonas et al. 2005). Dams and other barriers also represent a significantimpairment to many Great Lakes fish species. Migratory species, such as American Eel and Lake Sturgeon, are severely imperiled throughout most or all of the Great Lakes basin. Furthermore, dams in the Great Lakes basin are demonstrated to have a negative eff ect on fish species richness, particularly in small watersheds (Harford and McLaughlin 2007). However, dams are also an eff ective barrier for upstream migration of spawning Sea Lamprey (Harford and McLaughlin 2007; Lavis et al. 2003) and restrict the downstream movement of contaminated or 124 Brian M. Roth et al. nutrient-rich sediments into the lakes (Hart et al. 2002; Stanley and Doyle 2002). Managers must therefore carefully consider the positive benefits of increased stream connectivity with options for reducing down- stream sediment transport that will minimize unintended negative consequences for fish populations and fish consumers (Freeman et al. 2002).

Introduced Species

Th e Great Lakes off er one of the most striking examples of aquatic ecosystems altered by introduced species. More than one hundred eighty species have been documented introduced into the Great Lakes through various vectors (Ricciardi 2006). Th e construction of the Erie and Welland canals, the St. Lawrence Seaway, the Chicago Sanitary and Ship Canal, and the locks at Sault Ste. Marie have allowed several species access to the Great Lakes either directly through dispersal or indirectly by shipping ballast. Other species have been introduced intentionally through stocking, particularly salmonids, or accidentally through other means such as angler’s bait-buckets and the aquarium trade. A detailed account of introduced species in the Great Lakes can be found in Mandrak and Cudmore (2012). Th e number of species introduced into the Great Lakes continues to grow. Including reported and failed introductions, seventy-two fish and three decapod species have been introduced from outside the basin into at least one Great Lake basin; thirty-four of these species are now established in at least one Great Lake (table 5). Only nine of these species originated outside of North America. Twenty species originated from North American watersheds outside of the Great Lakes basin (table 5). Lakes Michigan and Erie support more species from North American basins than any other lake (thirteen and fourteen, respectively) most likely because of their southern location and proximity to donor basins (i.e., the Mississippi River basin). Vectors of species introduction are diverse. Some species, particularly salmonids, have been introduced intentionally to diversify and increase the commercial or recreational fishery and/or as a form of biological control for other problematic invasive species such as Alewife (Mills 1993). Others have been introduced as a result of the aquarium trade, such as the Goldfish and Oriental Weatherfish, or as baitfish (Rudd). Many species introduced into the Great Lakes are either a direct or an indirect consequence of the shipping industry (Holeck et al. 2004; Mills 1993; Ricciardi 2006). Ships entering the Great Lakes have introduced several species directly by exchanging ballast water, including the Round Goby, Tubenose Goby, and Ruff e. Th e canals built to circumvent natural barriers to shipping have also allowed several harmful species to expand their range from the Atlantic Coastal drainage, including Sea Lamprey, Alewife, Skipjack Herring, and White Perch. Similarly, connections to the Mississippi River have allowed species, such as the Yellow Bass, Orangespotted Sunfish, and perhaps the Red Shiner, to enter the Great Lakes basin.

TABLE 5. Summary of the Origins and Dispersal of Species Introduced into the Great Lakes

CATEGORY SUPERIOR MICHIGAN HURON ERIE ONTARIO TOTAL Total established 19 22 21 22 14 34 From inside basin 4 2 2 2 0 5 From outside basin 9 13 11 15 9 20 From outside North America 6 7 7 6 5 9 FISHES AND DECAPOD CRUSTACEANS 125

Th ree decapods represent prominent introductions into the Great Lakes: Rusty Crayfish, Red Swamp Crayfish, and Chinese Mitten Crab. Th e Rusty Crayfish likely dispersed from streams into the Great Lakes or were a bait-bucket introduction. Rusty crayfish is a notorious species introduced into every state and province in the Great Lakes basin (Lodge et al. 1985, 2000). Although most knowledge of this species as an invader derives from inland lakes and streams in the Mississippi and Great Lakes basin, dense populations of rusty crayfish could have profound eff ects on macrophytes in Great Lakes estuaries and wetlands, given their propensity to wreak similar havoc in smaller lakes (Lodge and Lorman 1987; Wilson et al. 2004). In contrast to rusty crayfish, the Red Swamp Crayfish is likely an intentional release from the aquarium trade (Simon and Th oma 2006; Th oma and Jezerinac 2000). Th e Red Swamp Crayfish is now common in the Sandusky Bay area of Lake Erie (Th oma and Jezerinac 2000). Th e Chinese Mitten Crab was first found in the Great Lakes in 1965 and likely arrived a stowaway in shipping ballast from Europe (Nepszy and Leach 1973). Th e Chinese Mitten Crab requires saltwater to complete its life cycle, so it is unlikely to establish a reproducing population within the Great Lakes, with the possible exception of Lake Ontario (de Lafontaine 2005). Nonetheless, the Chinese mitten crab has been collected in all the Great Lakes, except Lake Michigan (Benson and Fuller 2009).

Pollution

The influence of various forms of pollution on the fishes and decapods of the Great Lakes remains contentious. Although the vast quantities of pollutants were discharged into the Great Lakes, primarily from the 1800s until the 1960s (Christie 1974), partitioning toxicological eff ects on fish populations from those of other threats are difficult or impossible. Certainly, the Great Lakes were subjected to increased point-source and non-point-source pollution through industrial waste, increased fertilizer usage, the use of phosphate-based laundry detergent, and urbanization (Brazner et al. 2007). However, the eff ects of dioxins, DDT, PCBs, mercury, and other chemical contaminants are likely more subtle and likely pertain to the development and performance of various species (Murphy et al. 2005). Th e influence of these contaminants on fish populations remain under investigation. For more on this topic, see Murphy and Bhavsar (2012).

The Future of the Great Lakes Fish Community

Imminent Invasions

Over the last twenty years or so, several invasive fish species have established naturalized populations in watersheds adjacent to or near the Great Lakes drainage. Th ree notable species that pose imminent threats to Great Lakes ecosystems, given their eff ect in non-native waters, include the Northern Snakehead and Th e Northern Snakehead was previously introduced into Lake Michigan but apparently failed to establish a naturalized population. An angler caught a single Northern Snakehead in Chicago’s Burnham Harbor in 2004, but subsequent eff orts by biologists failed to capture any additional specimens. No other specimens have been discovered in Lake Michigan or surrounding waterbodies since this initial find, indicating the probable release of a fish from a live fish market or aquarium. However, Northern Snakehead have been found in Arkansas, New York, Pennsylvania, and Wisconsin and have established a naturalized population 126 Brian M. Roth et al. in Arkansas and the Potomac River in Maryland (Odenkirk and Owens 2007). Th e Northern Snakehead is a temperate species, and the entire Great Lakes basin has a suitable climate for a naturalized population (Herborg et al. 2007). Northern Snakehead prefer heavily vegetated habitat, which is abundant in wetlands, drowned river mouths, and the tributaries of the Great Lakes (Cudmore and Mandrak 2005). If this species becomes established in the Great Lakes basin, the potential for severe negative eff ects on the Great Lakes food web is high, given its piscivorous nature (Cudmore and Mandrak 2005). In response, all jurisdictions that surround the Great Lakes currently prohibit the sale, purchase, and transport of live Snakeheads. Asian carps also represent a significant threat tothe Great Lakes. Like the Northern Snakehead, Asian carps are temperate species and the entire Great Lakes basin is suitable for naturalization (Herborg et al. 2007). Both the Silver and Bighead carps have established naturalized populations in the middle Illinois River, and a single Bighead Carp was recently collected a mere 10 kilometers from Lake Michigan in Lake Calumet, Illinois. Asian carp DNA has also been found in Lake Michigan proper, suggesting they may already be in Lake Michigan. Bighead and Silver carps were first collected in the La Grange Reach of the Illinois River in 1995 and 1998, respectively. Since 2000, commercial catches of Bighead and Silver carps in the La Grange Reach have increased substantially, often exceeding 11,000 kilograms per day (Chick and Pegg 2001; Irons et al. 2007). Asian carps can move considerable distances over short periods of time (e.g., during flood pulses) and are expanding rapidly into upstream portions of the Illinois River. Mean maximum movement rates for Bighead and Silver carps were 6.8 and 10.6 km/day, respectively (Degrandchamp et al. 2008). Upstream movements of Asian carps and barriers to movement (i.e., locks and dams) have resulted in a distinct longitudinal gradient within the Illinois River, such that current abundances of Asian carps are lowest in the most upstream reaches and highest in the two lower reaches. However, abundances appear to be increasing in all reaches of the Illinois River. Th ese species prefer productive habitat that is available nearshore, in tributaries, and in drowned river mouths of the Great Lakes (Mandrak and Cudmore 2004). If one or more of the Asian carps become established in the Great Lakes basin, the potential for severe negative eff ects on the Great Lakes food web is high (Mandrak and Cudmore 2004). Th e sheer proximity of confirmed specimens to the Great Lakes warrants their inclusion in this chapter, and, if DNA evidence is correct, then Asian carps may already be present in Lake Michigan.

Future Prospects for Great Lakes Fish and Decapods

Th e fish and decapod communities of the Great Lakes is beset by a number of current and emerging stressors. Specific examples of stressors that could cause further community change include global climate change, non-native species, contaminants, eutrophication, habitat alteration, and fishing. Alone, each of these stressors is likely to cause changes in the Great Lakes fish community. For example, introductions of non-native species will likely continue. Kolar and Lodge (2002) listed a total of twenty-two species from the Caspian Sea region alone that could invade the Great Lakes, with five species (Tyulka Clupeonella cultriventris, Eurasian Minnow Phoxinus phoxinus, Black Sea Silverside Aphanius boyeri, European Perch Perca fluviatilis, and Monkey Goby Neogobius fluviatilis) likely to become nuisance species. Fishing and harvest management will likely also act on the Great Lakes food web, as they are inextricably linked (Kitchell and Hewitt 1987; Stewart and Ibarra 1991). Some of these stressors are likely to act synergistically, such that positive feedbacks may emerge. For example, Mandrak (1989) predicted based on discriminant analysis that global climate change could lead FISHES AND DECAPOD CRUSTACEANS 127 at least nineteen fish species to invade the Great Lakes basin from the Mississippi or Atlantic Coastal basins. Th e expansion of thermal habitat available to cool- and warm-water fishes opens previously untenable locations (e.g., much of Lake Superior) to colonization and establishment (Chu et al. 2005; Mandrak 1989; Sharma et al. 2007). Many crayfish species, currently limited by cold temperatures, are likely to migrate northward. Th e introduction of non-native species could also lead to dramatic changes in habitat, as is the case with Zebra (Dreissena polymorpha) and Quagga (D. rostriformis) mussels in open water areas of the Great Lakes or Eurasian Watermilfoil (Myriophyllum spicatum) and Purple Loosetrife (Lythrum salicaria) in wetlands and nearshore areas. Furthermore, global climate change is predicted to alter the hydrological cycle, with more precipitation in more northern regions of the basin, less precipitation in southern regions, and increased temperatures throughout (Magnuson et al. 1997; Mortsch and Quinn 1996). Changes in temperature and precipitation would likely aff ect available habitat for native species in Great Lake tributaries (McBean and Motiee 2008; Smith 1991). Additionally, warm-water crayfishes (e.g., Red Swamp Crayfish) could expand their range, which could lead to serious repercussions on aquatic macrophytes and the fishes that depend on them (Lodge and Lorman 1987; Lodge et al. 2000; Wilson et al. 2004). Th e combined eff ects of these stressors is still under evaluation, but integrated approaches that link population dynamics with climate eff ects are avenues for increased understanding of complex interactions (Jones et al. 2006). Th e history of fishing in the Great Lakes reveals a strong influence on the fish community of the Great Lakes. Th e sustainability of current fisheries depends on the ability of the ecosystem to support fish recruit- ment and productivity. Both recruitment and productivity are an emergent property of ecological condi- tions that can be altered by multiple stressors. Toxicological stress can initiate early mortality syndrome (EMS) in salmonids that consume prey high in thiaminase (e.g., Alewife or Rainbow Smelt) that, in turn, lead to thiamine deficiencies in embryos and larvae (Honeyfield et al. 2005). Other contaminants, such as dioxins, DDT, and PCBs, can also lead to sub-lethal and lethal eff ects on fish fitness and can severely impair reproduction of important predatory species, such as Lake Trout (Tillitt et al. 2008). Although contaminant concentrations are declining in most locations around the Great Lakes, climate change and the resulting drop in water levels could initiate re-suspension of toxins buried in sediments through dredging or from wind-borne sediments (Magnuson et al. 1997). Even if this scenario is speculative in terms of its eff ect on fish populations, increased human consumption advisories will likely result.

Summary

Change is a consistent theme of the fish and decapod communities of the Great Lakes basin. eTh retreat of the glaciers at the end of the Pleistocene allowed fishes and decapods to colonize the lakes and their watersheds. In modern times, changes in species composition in large part occurs through introductions. Various fish and decapod species were, in some cases, introduced intentionally and others accidentally. Th e shipping industry remains an important vector of non-native species in the Great Lakes (Holeck et al. 2004). Several fish species that derive from adjacent watersheds and overseas are predicted to invade the Great Lakes in the near future. Current eff ects of introduced species on Great Lakes ecosystems reinforce the need for management policies that place emphasis on preventing new invasions. If new introductions are inevitable, mitigation strategies that emphasize early detection and rapid responses are crucial to minimize potential negative eff ects. 128 Brian M. Roth et al.

We found a paucity (relative to fishes) of information on the distribution and zoogeography of decapods in the Great Lakes watershed. Th is lack of information is likely a result of the minimal direct economic importance (e.g., commercial and aquaculture fisheries) of these species in the Great Lakes basin. However, decapod crustaceans, and crayfishes in particular, have been labeled “keystone species” for their role in littoral and stream communities (Lodge and Hill 1994; Lodge et al. 1994; Momot 1995). Recent invasions and the discovery of new crayfish species in the Great Lakes basin (Th oma and Jezerinac 2000) highlight the need to characterize the distribution of decapod species, particularly in the Lake Huron, Ontario, and Superior basins. Th is need is exacerbated by global climate change predicted to expand the range of warm-water species northward (Magnuson et al. 1997; Sharma et al. 2007). Th e Great Lakes remain a unique ecosystem whose native fish community remains largely intact, despite stressors. Conserving fish resources remains adaunting challenge for managers in a world in which biological homogenization is increasingly common (Olden and Rooney 2006). Additional challenges lie in adapting to synergistic interactions between species introductions, habitat degradation, eutrophication, fishing, and global climate change. Further research, as well as prudent and cautious management, are necessary requirements to sustain the Great Lakes in a state resilient to these challenges.

Acknowledgments

We would like to thank the large number of individuals who shared their knowledge of the fish and crayfish communities with us. Th is chapter would not be possible without their help and willingness to share their experiences. Roger Th oma, David Lodge, the Canadian Museum of Nature, the Royal Ontario Museum, and the Ontario Federation of Anglers and Hunters were particularly helpful with decapod crustaceans. Becky Cudmore, Solomon David, Brian Gunderman, Scott Hanshue, Charles Madenjian, and Jay Wesley all reviewed the checklist of fishes in the Great Lakes andhelped clarify several disputes on distributions of various species. We would also like to thank Jana Lantry, Lars Rudstam, and Maureen Walsh for helping resolve questions regarding the existence and status of several fishes in the Lake Ontario basin.

REFERENCES

Bailey, R.M., W.C. Latta, and G.R. Smith. 2004. An atlas er.usgs.gov/queries/FactSheet.asp?speciesID=182, of Michigan fishes with keys and illustrations for revised 7/20/2007. their identification. Museum of Zoology, University Beeton, A. M. 1965. Eutrophication of the St. Lawrence of Michigan Miscellaneous Publication 192, Ann Great Lakes. Limnology and Oceanography Arbor, MI. 10:240–254. Bailey, R.M. and G.R. Smith. 1981. Origin and geography Bhagat, Y., J.J.H. Ciborowski, L.B. Johnson, D.G. Uzarski, of the fish fauna of the Laurentian Great Lakes T.M. Burton, S.T.A. Timmermans, and M.J. Cooper. Basin. Canadian Journal of Fisheries and Aquatic 2007. Testing a fish index of biotic integrity tor Sciences 38(12):1539–1561. responses to diff erent stressors in Great Lakes Becker, G.C. 1983. Fishes of Wisconsin. University of coastal wetlands. Journal of Great Lakes Research Wisconsin Press, London, England. 33:224–235. Benson, A.J. and P. Fuller. 2009. USGS nonindigenous Brazner, J.C. 1997. Regional, habitat, and human devel- aquatic species database, Gainesville, FL. http://nas. opment influences on coastal wetland and beach FISHES AND DECAPOD CRUSTACEANS 129

fish assemblages in Green Bay, Lake Michigan. Chu, C., N.E. Mandrak, and C.K. Minns. 2005. Potential Journal of Great Lakes Research 23(1):36–51. impacts of climate change on the distributions Brazner, J.C., N.P. Danz, A.S. Trebitz, G.J. Niemi, R.R. of several common and rare freshwater fishes in Regal, T. Hollenhorst, G.E. Host, E.D. Reavie, Canada. Diversity and Distributions 11(4):299–310. T.N. Brown, J.M. Hanowski, C.A. Johnston, L.B. Claramunt, R., J. Jonas, J.D. Fitzsimons, and J.E. Johnson, R.W. Howe, and J.J.H. Ciborowski. 2007. Marsden. 2005. Influences of spawning habitat Responsiveness of Great Lakes wetland indicators characteristics and interstitial predators on lake to human disturbances at multiple spatial scales: trout egg deposition and mortality. Transactions of a multi-assemblage assessment. Journal of Great the American Fisheries Society 134:1048–1057. Lakes Research 33:42–66. Cleland, C.E. 1982. Th e inland shore fishery of the north- Bridgeman, T., D. Schloesser, and A. Krause. 2006. ern Great Lakes: its development and importance in Recruitment of Hexagenia mayfly nymphs in west- prehistory. American Antiquity 47(4):761–784. ern Lake Erie linked to environmental variability. Coon, T.G. 1994. Ichthyofauna of the Great Lakes basin. Ecological Applications 16(2):601–611. Pages 55–71 in W.W. Taylor and C.P. Ferreri (eds.). Bronte, C., M. P. Ebener, D. R. Schreiner, D. S. DeVault, Great Lakes fisheries policy and management: a M. M. Petzold, D. A. Jensen, C. Richards, and S. binational perspective. Michigan State University J. Lozano. 2003. Fish community change in Lake Press, East Lansing, MI. Superior, 1970-2000. Canadian Journal of Fisheries COSEWIC. 2005. COSEWIC assessment and update and Aquatic Sciences 60:1552–1574. status report on the Lake Ontario kiyi Coregonus Brown, E.H., R.W. Rybicki, and R.J. Poff . 1985. Popula- kiyi orientalis and Upper Great Lakes kiyi Coregonus tion dynamics and interagency management of kiyi kiyi in Canada. Committee on the Status of the bloater (Coregonus hoyi) in Lake Michigan, Endangered Wildlife in Canada, Ottawa, ON. 1967–1982. Great Lakes Fishery Commission —. 2007. Update COSEWIC status report on Black Technical Report 23 Buff alo, Ictiobus niger. Committee on the Status of Carrick, H., J. Moon, and B. Gaylord. 2005. Phytoplank- Endangered Wildlife in Canada, Ottawa, ON. www. ton dynamics and hypoxia in Lake Erie: a hypothesis cosewic.gc.ca. concerning benthic-pelagic coupling in the central Crandall, K. and A. Templeton. 1999. Th e zoogeography basin. Journal of Great Lakes Research 31(Supple- and centers of origin of the crayfish subgenus ment 2):111–124. Procericambarus (Decapoda: Cambaridae). Evolu- Chapleau, F. and G. Pageau. 1982. Morphological tion 53(1):123–134. diff erentiation of Etheostoma olmstedi and E. Creaser, E.P. 1931. Th e Michigan decapod crustaceans. nigrum (Pisces: Percidae) in Canada. Copeia Papers of the Michigan Academy of Science, Arts, 1985(4):855–865. and Letters 13:257–276. Charlton, M.N. and J. Milne. 2004. Review of thirty Crossman, E.J. and D.E. McAllister. 1986. Zoogeography years of change in Lake Erie water quality. NWRI of freshwater fishes of the Hudson Bay drainage, Contribution No. 04-167. Ungava Bay and the Arctic Archipelago. Pages Chick, J.H. and M.A. Pegg. 2001. Invasive carp in the 53–104 in C.H. Hocutt and E.O. Wiley (eds.). Th e Mississippi River Basin. Science 292(5525):2250– zoogeography of North American freshwater fishes. 2251. John Wiley and Sons, New York, NY. Christie, W.J. 1974. Changes in fish species composition Cudmore, B. and N.E. Mandrak. 2005. Risk assessment of the Great Lakes. Journal of the Fisheries Research of northern snakehead (Channa argus) in Canada. Board of Canada 31(5):827–854. Canadian Science Advisory Secretariat, Fisheries 130 Brian M. Roth et al.

and Oceans Canada, Ottawa, Ontario Research Special Publication 05-01. Document, 2006/075, Ottawa, ON. France, R. 1992. Th e North American latutudinal gradi- Cudmore-Vokey, B. and E.J. Crossman. 2000. Checklists ent in species richness and geographical range of of the fish fauna of the Laurential Great Lakes and freshwater crayfish and amphipods. The American their connecting channels. Canadian Manuscript Naturalist 139(2):342–354. Report of Fisheries and Aquatic Sciences 2550. Fraser, B.A., N.E. Mandrak, and R.L. McLaughlin. 2005. Dahl, T.E. 1990. Wetlands losses in the United States Lack of morphological diff erentiation in eastern 1780s to 1980s. U.S. Fish and Wildlife Service, (Rhinichthys atratulus) and western (Rhinichthys Washington, DC. obtusus) blacknose dace in Canada. Canadian de Lafontaine, Y. 2005. First record of the Chinese Journal of Zoology-Revue Canadienne De Zoologie mitten crab (Eriocheir sinensis) in the St. Lawrence 83(11):1502–1509. River, Canada. Journal of Great Lakes Research Freeman, R., W. Bowerman, T. Grubb, A. Bath, G. 31(3):367–370. Dawson, K. Ennis, and J. Giesy. 2002. Opening rivers Degrandchamp, K.L., J.E. Garvey, and R.E. Colombo. to TROJAN fish: the ecological dilemma of dam 2008. Movement and habitat selection by invasive removal. Conservation in Practice 3:35–40. Asian carps in a large river. Transactions of the Fuller, P. 2009. Ictiobus bubalus. USGS nonindigenous American Fisheries Society 137(1):45–56. aquatic species database. http://nas.er.usgs.gov/ Docker, M., N.E. Mandrak, D. Heath, and K. Scribner. queries/FactSheet.asp?speciesID=361 Revised 2005. Genetic markers to distinguish and quantify 4/18/2006. the level of gene flow between northern brook and Grande, T., H. Laten, and J. Lopez. 2004. Phylogenetic silver lampreys. Final Completion Report, Great relationships of extant esocid species (Teleostei: Lakes Fishery Commission, Ann Arbor, MI. Salmoniformes) based on morphological and Dodge, D. and R. Kavetsky. 1994. Aquatic habitat and molecular characters. Copeia 2004(4):743–757. wetlands of the Great Lakes. State of the Great Lakes Hansen, M. J. 1994. Th e state of Lake Superior in 1992. Ecosystem Working Paper. EPA 905-D-94-001C. U.S. Great Lakes Fish. Comm. Spec. Pub. 94-1. Environmental Protection Agency, Washington, DC Harford, W.J. and R.L. McLaughlin. 2007. Understanding and Environment Canada, Ottawa, ON. uncertainty in the eff ect of low-head dams on fishes Dryer, W., L. Erkkila, and C. Tetzloff . 1965. Food of the of Great Lakes tributaries. Ecological Applications lake trout in Lake Superior. Transactions of the 17(6):1783–1796. American Fisheries Society 94:169–176. Hart, D.D., T.E. Johnson, K.L. Bushaw-Newton, R.J. Edsall, T. and G.W. Kennedy. 1995. Availability of lake Horwitz, A.T. Bednarek, D.F. Charles, D.A. Kreeger, trout reproductive habitat in the Great Lakes. and D.J. Velinsky. 2002. Dam removal: challenges Journal of Great Lakes Research 21(Supplement and opportunities for ecological research and river 1):290–301. restoration. BioScience 52(8):669–381. Edwards, W., J. Conroy, and D. Culver. 2005. Hypolim- Herborg, L., C.L. Jerde, D.M. Lodge, G.M. Ruiz, and netic oxygen depletion dynamics in the central H.J. MacIsaac. 2007. Predicting invasion risk basin of Lake Erie. Journal of Great Lakes Research using measures of introduction eff ort and envi- 31(Supplement 2):262–271. ronmental niche models. Ecological Applications Fleischer, G.W., C.P. Madenjian, R.F. Elliott, and M.L. 17(3):663–674. Toneys. 2005. Planktivores. Pages 16–20 in M.E. Hobbs, H.H. 1974. A checklist of the North and Middle Holey and T.N. Trudeau (eds.). Th e state of Lake American crayfishes (Decapodae: Astacidae and Michigan in 2000. Great Lakes Fishery Commission Cambaridae). Smithsonian Institution Press, FISHES AND DECAPOD CRUSTACEANS 131

Washington, DC. Canandian Journal of Fisheries and Aquatic Sci- Hobbs, H.H.I. and J.P. Jass. 1988. Th e crayfishes and ences 62:2254–2264. shrimps of Wisconsin. Wisconsin Public Museum, Jones, M.L., B. Schuter, Y. Zhao, and J. Stockwell. 2006. Milwaukee, WI. Forecasting eff ects of climate change on Great Hoff , M.H. and T.N. Todd. 2004. Status of the shortjaw Lakes fisheries: models that link habitat supply to cisto (Coregonus zenithicus) in Lake Superior. population dynamics can help. Canandian Journal Annales Zoologici Fennici 41:147–154. of Fisheries and Aquatic Sciences 63:457–468. Holeck, K.T., E.L. Mills, H.J. MacIsaac, M.R. Dochoda, Kelso, J.R.M., R.J. Steedman, and S. Stoddart. 1996. His- R.I. Colautti, and A. Ricciardi. 2004. Bridging torical causes of change in Great Lakes fish stocks troubled waters: biological invasions, transoceanic and the implications for ecosystem rehabilitation. shipping, and the Laurentian Great Lakes. BioSci- Canadian Journal of Fisheries and Aquatic Sciences ence 54(10):919–929. 53(Supplement 1):10–19. Honeyfield, D. C., J. P. Hinterkopf, J. D. Fitzsimons, D. E. Kitchell, J.F. and S.W. Hewitt. 1987. Forecasting forage Tillitt, J. L. Zajicek, and S. B. Brown. 2005. Develop- demand and yield of sterile chinook salmon ment of thiamine deficiencies and early mortality (Oncorhynchus tshawytscha) in Lake Michigan. syndrome in lake trout by feeding experimental and Canadian Journal of Fisheries and Aquatic Sciences feral fish diets containing thiaminase. Journal of 44(Supplement 2):384–389. Aquatic Animal Health 17:4–12. Koelz, W. 1929. Coregonid fishes of the Great Lakes. Hrabik, T. R., O. Jensen, S. Martell, C. J. Walters, and J. Bulletin of the US Bureau of Fisheries 43:297–643. F. Kitchell. 2006. Diel vertical migration in the Lake Kolar, C.S. and D.M. Lodge. 2002. Ecological predic- Superior pelagic community I. Changes in vertical tions and risk assessment for alien fishes in North migration of coregonids in response to varying America. Science 298:1233–1236. predation risk. Canadian Journal of Fisheries and Kott, E. and D. Fitzgerald. 2000. Comparative morphol- Aquatic Sciences 63:2286–2295. ogy and taxonomic status of the ghost shiner, Hubbs, C.L. and K.F. Lagler. 2004. Fishes of the Great Notropis buchanani, in Canada. Environmental Lakes Region. University of Michigan Press, Ann Biology of Fishes 59(4):385–392. Arbor, MI. Lantry, B., R. O’Gorman, M. Walsh, J. Casselman, J. Hubert, N., R. Hanner, E. Holm, N.E. Mandrak, E. Taylor, Hoyle, M. Keir, and J. Lantry. 2007. Reappearance of M. Burridge, D. Watkinson, P. Dumont, A. Curry, P. deepwater sculpin in Lake Ontario: resurgence or Bentzen, J. Zhang, J. April, and L. Bernatchez. 2008. last gasp of a doomed population? Journal of Great Identifying Canadian freshwater fishes through DNA Lakes Research 33(Supplement 1):34–45. barcodes. PLoS ONE 3(6):e2490. Lavis, D.S., A. Hallett, E.M. Koon, and T.C. McAuley. Irons, K., G. Sass, M. McClelland, and J. Staff ord. 2007. 2003. History of and advances in barriers as an Reduced condition factor of two native fish species alternative method to suppress sea lampreys in coincident with invasion of non-native Asian carps the Great Lakes. Journal of Great Lakes Research in the Illinois River, U.S.A. Is this evidence for 29(Supplement 1):362–372. competition and reduced fitness? Journal of Fish Lawrie, A. and J. Rahrer. 1973. Lake Superior: a case Biology 71(Supplement D):258–273. history of the lake and its fisheries. Great Lakes Jonas, J., R. Claramunt, J.D. Fitzsimons, J.E. Marsden, Fishery Commission, Technical Report 19. Great and B. Ellrott. 2005. Estimates of egg deposition Lakes Fishery Commission, Ann Arbor, MI. and eff ects of lake trout (Salvelinus namaycush) Lindsey, C.C. 1981. Stocks are chameleons—plasticity in egg predators in three regions of the Great Lakes. gill rakers of coregonid fishes. Canadian Journal of 132 Brian M. Roth et al.

Fisheries and Aquatic Sciences 38(12):1497–1506. Mandrak, N.E. 2009. Fish fauna of Lake Superior: past, Lodge, D.M. and A.M. Hill. 1994. Factors govern- present and future. Pages 645–663 in M. Munawar ing species composition, population size, and and I.F. Munawar (eds.). State of Lake Superior. productivity of cool-water crayfishes. Nordic Journal Ecovision World Monograph Series. of Freshwater Research 69:111–136. Mandrak, N.E. and E.J. Crossman. 1992. A checklist of Lodge, D.M. and J.G. Lorman. 1987. Reductions in Ontario freshwater fishes. Royal Ontario Museum, submersed macrophyte biomass and species Toronto, ON. richness by the crayfishOrconectes rusticus. Mandrak, N.E. and B. Cudmore. 2004. Risk assessment Canadian Journal of Fisheries and Aquatic Sciences for Asian carps in Canada. Canadian Science 44:591–597. Advisory Secretariat, Fisheries and Oceans Canada, Lodge, D.M., A.L. Beckel, and J.J. Magnuson. 1985. Lake- Ottawa, Ontario Research Document. 2004/103, Bottom Tyrant. Natural History 1985(8):33–37. Ottawa, ON. Lodge, D.M., M.W. Kershner, J.E. Aloi, and A.P. Covich. Mandrak, N.E. and B. Cudmore. 2012. Fish species 1994. Eff ects of an omnivorous crayfish Orconectes( at risk and non-native fishes in the Great Lakes rusticus) on a freshwater littoral food web. Ecology basin: past, present, and future. Pages •••–••• in 75(5):1265–1281. W.W. Taylor, A.J. Lynch, and N.J. Leonard (eds.). Lodge, D.M., C.A. Taylor, D.M. Holdich, and J. Skurdal. Great Lakes fisheries policy and management: a 2000. Nonidigenous crayfishes threaten North binational perspective (second edition). Michigan American freshwater biodiversity. Fisheries State University Press, East Lansing, MI. 25(8):7–19. Mandrak, N.E., M. Docker, and D. Heath. 2004. Native Madenjian, C.P., D.M. Warner, D.B. Bunnell, R. Ichthyomyzon lampreys of the Great Lakes: develop- Claramunt, and J.M. Dettmers. 2008. Status of ment of genetic markers and a morphological key planktivore populations. Pages 49-54 in D.F. Clapp to ammocoetes. Annual Report, Great Lakes Fishery and W. Horns (eds.). Th e state of Lake Michigan Commission, Ann Arbor, MI. in 2005. Great Lakes Fishery Commission Special Markham, J.L. 2009. Past and present salmonid Publication 08-02, Ann Arbor, MI. community of Lake Erie. Pages 41–50 in J.T. Tyson, Magnuson, J.J., G.M. Capelli, J.G. Lorman, and R.A. R.A. Stein, and J.M. Dettmers (eds.). Th e state of Stein. 1975. Consideration of crayfish for macro- Lake Erie in 2004. Great Lakes Fishery Commission phyte control. Pages 66–74 in P.L. Brezonik and Special Publication 09-02. J.C. Fox (eds.). Th e Proceedings of a Symposium McBean, E. and H. Motiee. 2008. Assessment of impact on Water Quality Management through Biological of climate change on water resources: a long term Control. Report No. ENV 07-75-1, Gainesville, FL. analysis of the Great Lakes of North America. Hy- Magnuson, J.J., K.E. Webster, R.A. Assel, C.J. Bowser, drology and Earth System Sciences 12(1):239–255. P.J. Dillon, J.G. Eaton, H.E. Evans, E.J. Fee, R.I. Mills, E.L. 1993. Exotic species in the Great Lakes: a Hall, L.R. Mortsch, D.W. Schindler, and F.H. history of biotic crises and anthropogenic introduc- Quinn. 1997. Potential eff ects of climate changes tions. Journal of Great Lakes Research 19(1):1–54. on aquatic systems: Laurentian Great Lakes and Mills, E.L., J. Casselman, R. Dermott, J.D. Fitzsimons, Precambrian Shield Region. Hydrological Processes G. Gal, K.T. Holeck, J. Hoyle, O.E. Johannsson, B. 11(8):825–871. Lantry, J. Makarewicz, E. Millard, I. Munawar, M. Mandrak, N.E. 1989. Potential invasion of the Great Munawar, R. O’Gorman, R. Owens, L.G. Rudstam, Lakes by fish species associated with climatic warm- T. Schaner, and T.J. Stewart. 2003. Lake Ontario: ing. Journal of Great Lakes Research 15(2):306–316. food web dynamics in a changing ecosystem FISHES AND DECAPOD CRUSTACEANS 133

(1970–2000). Canadian Journal of Fisheries and Ichthyomyzon ammocoetes using meristic, mor- Aquatic Sciences 60:471–490. phological, pigmentation, and gonad analyses. Mills, E.L., J. Casselman, R. Dermott, J.D. Fitzsimons, Canadian Journal of Zoology-Revue Canadienne De G. Gal, K.T. Holeck, J. Hoyle, O.E. Johannsson, B. Zoologie 85(4):549–560. Lantry, J. Makarewicz, E. Millard, I. Munawar, M. Nelson, J., E.J. Crossman, H. Espinosa-Perez, L. Findley, Munawar, R. O’Gorman, R. Owens, L.G. Rudstam, C. Gilbert, R. Lea, and A.H. Williams. 2004. A list of T. Schaner, and T.J. Stewart. 2005. A synthesis of scientific and common names of fishes from the ecological and fish community changes in Lake United States and Canada, Sixth Edition. American Ontario, 1970–2000. Great Lakes Fishery Commis- Fisheries Society Special Publication 29, Bethesda, sion Technical Report 67. MD. Minns, C.K., V.W. Cairns, R.G. Randall, and J.E. Moore. Nepszy, S.J. and J.H. Leach 1973. First records of the 1994. An index of biotic integrity (IBI) for fish as- Chinese mitten crab, Eriocheir sinensis, (Crustacea: semblages in the littoral zone of Great Lakes areas of Brachyura) from North America. Journal of the Fish- concern. Canadian Journal of Fisheries and Aquatic eries Research Board of Canada 30(12):1909–1910. Sciences 51(8):1804–1822. Odenkirk, J. and S. Owens. 2007. Expansion of a north- Mohr, L.C. and M.P. Ebener. 2005. Th e coregonine com- ern snakehead population in the Potomac River munity. Special Publication. Great Lakes Fishery system. Transactions of the American Fisheries Commission 05-02:69–76. Society 136:1633–1639. Momot, W.T. 1995. Redefining the role of crayfish in Olden, J. and T. Rooney. 2006. On defining and quantify- aquatic ecosystems. Reviews in Fisheries Science ing biotic homogenization. Global Ecology and 3(1):33–63. Biogeography 15:113–120. Mortsch, L.D. and F.H. Quinn. 1996. Climate change Owens, R., R. O’Gorman, E.L. Mills, L.G. Rudstam, J. scenarios for Great Lakes basin ecosystem studies. Hasse, B. Kulik, and D.R. MacNeill. 1998. Blueback Limnology and Oceanography 41(5):903–911. herring (Alosa aestivalis) in Lake Ontario: first Murdoch, M.H. and P.D.N. Hebert. 1994. Mitochondrial record, entry route, and colonization potential. DNA diversity of brown bullhead from contami- Journal of Great Lakes Research 24(3):723–730. nated and relatively pristine sites in the Great Lakes. Page, L.M. 1985. Th e crayfishes and shrimps (Decapoda) Environmental Toxicology 13(8):1281–1289. of Illinois. Illinois Natural History Survey Bulletin Murphy, C.A., K.A. Rose, and P. Th omas. 2005. Modeling 33. vitellogenesis in female fish explosed to environ- Perry, W. L., D. M. Lodge, and G. A. Lamberti. 1997. mental stressors: predicting the eff ects of endocrine Impact of crayfish predation on exotic zebra mus- disturbance due to exposure to a PCB mixture and sels and native invertebrates in a lake-outlet stream. cadmium. Reproductive Toxicology 19:395–409. Canadian Journal of Fisheries and Aquatic Sciences Murphy, C.A., S.P. Bhavsar, and N. Ghandi. 2012. 54:120–125. Contaminants in Great Lakes fish: historical, Petzold, M.M. 2002. Distribution and abundance current and emerging concerns. Pages - in of chub species (Coregonus sp.) in eastern Lake W.W. Taylor, A.J. Lynch, and N.J. Leonard (eds.). Superior (2000–2001). Upper Great Lakes Manage- Great Lakes fisheries policy and management: a ment Unity, Ontario Ministry of Natural Resources binational perspective (second edition). Michigan Technical Report 1-2002. State University Press, East Lansing, MI. Pratt, T.C. and N.E. Mandrak. 2007. Abundance, Neave, F.B., N.E. Mandrak, M.F. Docker, and D.L. distribution and identification of the shortjaw Noakes. 2007. An attempt to diff erentiate sympatric cisco (Coregonus zenthicus) in the proposed Lake 134 Brian M. Roth et al.

Superior marine protected area. Canadian Technical of the Laurentian Great Lakes: health, habitat, and Report of Fisheries and Aquatic Sciences 2697. indicators. AuthorHouse, Bloomington, IN. Pritchard, A.L. 1931. Taxonomic and life history studies Sly, P. and D. Evans. 1996. Suitability of habitat for of the ciscoes of Lake Ontario. University of Toronto spawning lake trout. Journal of Aquatic Ecosystem Studies in Biology Series 35. Publication of the Health 5:153–175. Ontario Fisheries Research Laboratory 41. Smith, J.B. 1991. Th e potential impacts of climate Randall, R.G., C.K. Minns, V.W. Cairns, and J.E. Moore. change on the Great Lakes. Bulletin of the American 1996. Th e relationship between an index of fish Meteorological Society 72(1):21–28. production and submerged macrophytes and other Smith, S.H. 1964. Status of the deepwater cisco popula- habitat features at three littoral areas in the Great tion of Lake Michigan. Transactions of the American Lakes. Canadian Journal of Fisheries and Aquatic Fisheries Society 93:155–163. Sciences 53(Supplement 1):35–44. Stanley, E.H. and M.W. Doyle. 2002. A geomorphic Ricciardi, A. 2006. Patterns of invasion of the Laurentian perspective on nutrient retention following dam Great Lakes in relation to changes in vector activity. removal. BioScience 52(8):693–701. Diversity and Distributions 12:425–433. Stewart, D.J. and M. Ibarra. 1991. Predation and Roseman, E., D. Jude, M. Raths, T. Coon, and W. Taylor. production by salmonine fishes in Lake Michigan, 1998. Occurrence of the deepwater sculpin (Myoxo- 1978–1888. Canadian Journal of Fisheries and cephalus thompsoni) in western Lake Erie. Journal of Aquatic Sciences 48:909–922. Great Lakes Research 24(2):479–483. Stone, U.B. 1947. A study of the deep-water cisco fishery Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Lake Ontario with particular reference to the of Canada. Fisheries Research Board of Canada bloater, Leucichthys hoyi (Gill). Transactions of the Bulletin 184. American Fisheries Society 74:230–249. Scott, W.B. and S.H. Smith. 1962. Th e occurrence of the Th oma, R. and R. Jezerinac. 2000. Ohio crayfish and longjaw cisco, Leucichthys alpenae, in Lake Erie. shrimp atlas. Ohio Biological Survey Miscellaneous Canadian Journal of Fisheries and Aquatic Sciences Contributions Number 7. 19:1013–1023. Tillitt, D., P. Cook, J. Giesy, W. Heideman, and R. Peter- Selgeby, J. 1982. Decline of lake herring (Coregonus son. 2008. Reproductive impairment of Great Lakes artedii) in Lake Superior: an analysis of the Wiscon- Lake Trout by Dioxin-like chemicals. Pages 819–875 sin herring fishery, 1936-78. Canadian Journal of in R. DiGiulio and D. Hinton (eds.). Th e toxicology Fisheries and Aquatic Sciences 39:554–563. of fishes. CRC Press, Boca Raton, FL. Selgeby, J., C.R. Bronte, and J. Slade. 1994. Forage Todd, T.N. 1985. Status of Great Lakes coregonines. species. Pages 48–57 in M.J. Hansen (ed.). Th e Great Lakes Fisheries Laboratory, Ann Arbor, MI. state of Lake Superior in 1992. Great Lakes Fishery Todd, T.N. and G.R. Smith. 1980. Diff erentiation in Core- Commission Special Publication 94-1. gonus zenithicus in Lake Superior. Canadian Journal Sharma, S., D.A. Jackson, C.K. Minns, and B.J. Shuter. of Fisheries and Aquatic Sciences 37(12):2228–2235. 2007. Will northern fish populations be in hot water Todd, T.N., G.R. Smith, and L.E. Cable. 1981. Environ- because of climate change? Global Change Biology mental and genetic contributions to morphological 13(10):2052–2064. diff erentiation in ciscoes (Coregoninae) of the Great Simon, T. and R. Th oma. 2006. Native and alien distribu- Lakes. Canadian Journal of Fisheries and Aquatic tion of crayfish assemblages in drowned river mouth Sciences 38(1):59–67. coastal wetlands of Lake Michigan. Pages 375–382 Trautman, M.B. 1981. Th e fishes of Ohio. Ohio State in T. Simon and P. Stewart (eds.). Coastal wetlands University Press, Columbus, OH. FISHES AND DECAPOD CRUSTACEANS 135

Turgeon, J. and L. Bernatchez. 2003. Reticulate evolution Ontario. Pages 51–57 in H.F. Allen, J.F. Reinwand, and phenotypic diversity in North American ciscoes, R.E. Ogawa, J.K. Hiltunen, and L. Wells (eds.). Coregonus spp. (Teleostei : Salmonidae): implica- Limnological survey of Lake Ontario, 1964. Great tions for the conservation of an evolutionary legacy. Lakes Fishery Commission Technical Report 14. Conservation Genetics 4(1):67–81. Williams, A.B., L.G. Abele, D.L. Felder, H.H. Hobbs Turgeon, J., A. Estoup, and L. Bernatchez. 1999. Species Jr., R.B. Manning, P.A. McLaughlin, and I. Perez flock in the North American Great Lakes: molecular Farfante. 1989. Common names of aquatic inverte- ecology of Lake Nipigon Ciscoes (Teleostei: Corego- brates from the United States and Canada: decapod nidae: Coregonus). Evolution 53(6):1857–1871. crustaceans. American Fisheries Society Special Tyson, J.T. and R.L. Knight. 2001. Response of yellow Publication 17, Bethesda, MD. perch to changes in the benthic invertebrate com- Wilson, C.C. and P.D.N. Hebert. 1996. Pylogeographic munity of western Lake Erie. Transactions of the origins of lake trout (Salvelinus namaycush) in American Fisheries Society 130:766–782. North America. Canadian Journal of Fisheries and Underhill, J.C. 1986. Th e fish fauna of the Laurentian Aquatic Sciences 53:2764–2775. Great Lakes, the St. Lawrence lowlands, Newfound- Wilson, C.C. and P.D.N. Hebert. 1998. Phylogeographic land and Laborador. Pages 105–136 in C.H. Hocutt and postglacial dispersal of Lake Trout (Salvelinus and E.O. Wiley (eds.). Th e zoogeography of North namaycush) in North American. Canadian Journal American freshwater fishes. John Wiley & Sons, Inc., of Fisheries and Aquatic Sciences 55:1010–1024. New York, NY. Wilson, K.A., J.J. Magnuson, D.M. Lodge, A.M. Hill, T.K. Van Oosten, J. 1937. Th e dispersal of smelt, Osmerus Kratz, W.M. Perry, and T.V. Willis. 2004. A long- mordax (Mitchill), in the Great Lakes Region. term rusty crayfish Orconectes( rusticus) invasion: Transactions of the American Fisheries Society dispersal patterns and community change in a north 66:160–171. temperate lake. Canadian Journal of Fisheries and Wells, L. 1969. Fishery survey of U.S. waters of Lake Aquatic Sciences 61: 2255–2266. View publication stats