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IchthyofaunaoftheGreatLakesBasin

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105

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

Brian M. Roth et al.106

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, and the 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

Brian M. Roth et al.108

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

Brian M. Roth et al.110

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

Brian M. Roth et al.112

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 routes have 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 P ROPOSED 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

Brian M. Roth et al.114

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.

Brian M. Roth et al.116

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

Brian M. Roth et al.118

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). Th e 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 Great Lakes 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

Brian M. Roth et al.120

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

Brian M. Roth et al.122

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 growth of 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 many lakes (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 significant impairment 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

Brian M. Roth et al.124

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

Brian M. Roth et al.126

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 to the 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. Th e 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.

Brian M. Roth et al.128

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 a daunting 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 and helped 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.

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