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Electronic Theses and Dissertations

2020

Fishes of the Dakotas

Kathryn Schlafke South Dakota State University

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Recommended Citation Schlafke, Kathryn, "Fishes of the Dakotas" (2020). Electronic Theses and Dissertations. 3942. https://openprairie.sdstate.edu/etd/3942

This Thesis - Open Access is brought to you for free and open access by Open PRAIRIE: Open Public Research Access Institutional Repository and Information Exchange. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of Open PRAIRIE: Open Public Research Access Institutional Repository and Information Exchange. For more information, please contact [email protected]. FISHES OF THE DAKOTAS

BY

KATHRYN SCHLAFKE

A thesis submitted in partial fulfillment of the requirements for the

Master of Science

Major in Wildlife and Fisheries Sciences

Specialization in Fisheries Science

South Dakota State University

2020

ii

THESIS ACCEPTANCE PAGE

Kathryn Schlafke

This thesis is approved as a creditable and independent investigation by a candidate for

the master’s degree and is acceptable for meeting the thesis requirements for this degree.

Acceptance of this does not imply that the conclusions reached by the candidate are

necessarily the conclusions of the major department.

Brian Graeb, Ph.D. Advisor Date

Michele R. Dudash Department Head Date

Dean, Graduate School Date iii

ACKNOWLEDGMENTS

I would first like to thank my advisors throughout this project, Dr. Katie

Bertrand and Dr. Brian Graeb for giving me the opportunity to work towards a graduate degree at South Dakota State University. Brian, thank you for always appearing to be calm in my states of panic, for always encouraging me, and for giving me the opportunities to express what I think when making decisions. I’m thankful for your continued help and support throughout this journey. I would also like to thank my committee members, Dr. Steven Chipps, Chelsey Pasbrig and Geno Adams for their time and support, as well as being patient with me. This project would also not be possible without the assistance of Matt Wagner and Nick Kludt. Matt, thank you for getting this project started to a point where I was able to jump in. I know it took an incredible amount of time to gather all the information for the distribution maps, create a dichotomous key, and to take your stunning photos. I appreciate everything you’ve done to get the project to where it is now, and for providing assistance remotely. Nick, thank you for your time and patience to help create the maps needed for this book.

Your wisdom in GIS will never cease to amaze me.

This project would not have been possible without the support of several South

Dakota Game, Fish and Parks Fisheries staff members across the state. I would like to express my sincere gratitude to the various members who contributed to edits on numerous species accounts for this book, as well as going out to sample for species throughout the duration of this project. I greatly appreciate the time and effort of every individual involved and cannot thank them enough for providing me with the information needed to complete this project. A sincere thank you is also extended out to iv

Chelsey Pasbrig for taking several hours of her time to edit numerous species accounts, introduction chapters, provide ideas and insight, and for continuing to support and check up on me throughout the entirety of this project.

A special thank you to all the friends I have made here at SDSU in the NRM department. I am forever grateful that we crossed paths and I hope we continue to do so in our futures. I am so thankful for the supportive community you have all contributed to, and I am not quite sure how I would have survived graduate school without each one of you. Thank you for the great conversations, always listening to me, and for all the fun times. I am also very grateful and appreciative of the support from the Natural

Resource Management office staff who have helped make not only my own, but everyone else’s lives a lot easier.

I would also like to extend a thank you to all of the many professionals and graduate students throughout my undergraduate career who encouraged me to pursue my education, and who guided and mentored me through opportunities to help develop skills and personal growth. Thank you for continuing to reach out throughout my entire graduate experience, and I am very thankful and appreciative of our relationships.

Last, but certainly not least, I owe a tremendous amount of thanks to my amazing family. Mom, Dad, William, and Myla, thank you for being so understanding, supportive, and encouraging throughout this entire process. Throughout all the twists, turns, and curveballs, I could always count on you to listen to me and pick me back up with a little (or a lot) of laughter. You guys are the absolute best, and I am so thankful to call you mine. I would not be where I am today without you, so thank you. v

This project was made possible and administered by the funding provided from

South Dakota Game, Fish and Parks and the Agricultural Experiment Station. A special thanks to SDSU and the Department of Natural Resource Management for providing the necessary resources.

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TABLE OF CONTENTS

LIST OF FIGURES…………………………………………………………………..xvii

ABSTRACT……………………………………………………………………...….xviii

CHAPTER 1: WATERS AND GEOLOGY OF NORTH DAKOTA AND SOUTH

DAKOTA………………………………………………………………………………..1

Level III Ecoregions…………………………………………………………….7

Lake Agassiz Plain………………………………………………………………7

Western Corn Belt Plains………………………………………………………..8

Northern Glaciated Plains…………………………………………………….....9

Northwestern Glaciated Plains…………………………………………………10

Northwestern Great Plains……………………………………………………..11

Nebraska Sand Hills…………………………………………………………....13

High Plains……………………………………………………………………..14

Middle Rockies……………………………………………………………...…15

Major River Basins of the Dakotas………………………………………….…16

Bad River Basin………………………………………………………..18

Belle Fourche River Basin ………………………………………...…..18

Big Sioux River Basin ………………………………………………...19

Cannonball River Basin ……………………………………………….19 vii

Cheyenne River Basin………………………………………………….19

Devils Lake River Basin…………………………………….…………20

Grand River Basin…………………………………………….………..21

Heart River Basin…………………………………………………..…..21

James River Basin………………………………………….…………..22

Knife River Basin……………………………………………………...22

Little Missouri River Basin…………………………………………….23

Minnesota River Basin…………………………………………………23

Missouri River Basin…………………………………………………..24

Garrison Dam…………………………………………………..25

Oahe Dam…………………………………………….………..26

Big Bend Dam ……………………………………..…………..26

Fort Randall Dam………………………………………..……..27

Gavins Point Dam………………………………………..…….27

Moreau River Basin……………………………………………………28

Niobrara River Basin…………………………………………….…….29

Red River of the North River Basin………………………..…………..29

Sheyenne River Basin………………………………………………….30 viii

Souris River Basin……………………………………………………..30

Vermillion River Basin…………………………………..…………….31

White River Basin……………………………………………….……..31

Literature Cited…………………………………………………….…………..32

CHAPTER 2: HISTORY OF ICHTHYOLOGY IN THE DAKOTAS………………..38

Prehistoric Fishes………………………………………………………………38

1800’s…………………………………………………………………………..38

Early 1900’s………………………………………………………………...….41

Post Reservoir Construction: 1950’s to Today…………………………...……43

Literature Cited……………………………………………………………..….45

CHAPTER 3: FAMILY PETROMYZONTIDAE……………………………………..46

Chestnut Lamprey, Ichthyomyzon castaneus………………………………..…48

Silver Lamprey, Ichthyomyzon unicuspis………………………………...……53

CHAPTER 4: FAMILY ACIPENSERIDAE…………………………………...…..…...58

Lake Sturgeon, Acipenser fulvescens………………………………………….…60

Pallid Sturgeon, Scaphirhynchus albus………………………………………..…66

Shovelnose Sturgeon, Scaphirhynchus platorynchus……………………………72

CHAPTER 5: FAMILY POLYODONTIDAE……………………………………….….77 ix

Paddlefish, Polyodon spathula…………………………………………….……..80

CHAPTER 6: FAMILY LEPISOSTEIDAE…………………………………….……..85

Longnose Gar, Lepisosteus osseus……………………………………….…….87

Shortnose Gar, Lepisosteus platostomus……………………………………….92

CHAPTER 7: FAMILY HIODONTIDAE………………………………...…………..97

Goldeye, Hiodon alosoides……………………………………………...……..99

Mooneye, Hiodon tergisus……………………………………………………103

CHAPTER 8: FAMILY ANGUILLIDAE……………………………………………108

American Eel, Anguilla rostrate……………………………………….……..110

CHAPTER 9: FAMILY CLUPEIDAE……………………………………………….117

Alewife, Alosa pseudoharengus……………………………………………...119

Gizzard Shad, Dorosoma cepedianum………………………………………..124

Skipjack Herring, Alosa chrysochloris…………………………...…………..130

Threadfin Shad, Dorosoma petenense………………………………………..134

CHAPTER 10: FAMILY CYPRINIDAE…………………………………………….139

Central Stoneroller, Campostoma anomalum………………………...………141

Largescale Stoneroller, Campostoma oligolepis…………………………..….146

Goldfish, Carassius auratus………………………………………………….151 x

Northern Redbelly Dace, Chrosomus eos……………………………...……..156

Southern Redbelly Dace, Chrosomus erythrogaster…………………………162

Finescale Dace, Chrosomus neogaeus………………………………………..167

Lake Chub, Couesius plumbeus………………………………………………172

Grass Carp, Ctenopharyngodon Idella………………………………...……..177

Red Shiner, Cyprinella lutrensis…………………………………………...…183

Spotfin Shiner, Cyprinells spiloptera……………………………………...…189

Common Carp, Cyprinus carpio………………………………...……………194

Western Silvery Minnow, Hybognathus argyritis………………………...….200

Brassy Minnow, Hybognathus hankinsoni…………………………….……..206

Plains Minnow, Hybognathus placitus……………………………………….211

Silver Carp, Hypophthalmichthys molitrix…………………………….……..217

Bighead Carp, Hypophthalmichthys nobilis………………………………….223

Common Shiner, Luxilus cornutus……………………………………...……229

Sturgeon Chub, Macrhybopsis gelida…………………………………….…..234

Shoal Chub, Macrhybopsis hyostoma………………………………….……..239

Sicklefin Chub, Macrhybopsis meeki………………………………….……..244

Silver Chub, Macrhybopsis storeriana………………………………...……..251 xi

Northern Pearl Dace, Margariscus nachtriebi………………………………..257

Hornyhead Chub, Nocomis biguttatis………………………………….……..263

Golden Shiner, Notemigonus crysoleucas……………………………..……..268

Pugnose Shiner, Notropis anogenus………………………………….………273

Emerald Shiner, Notropis atherinoides……………………………………….278

River Shiner, Notropis blennius……………………………………..………..283

Bigmouth Shiner, Notropis dorsalis………………………………...………..287

Blacknose Shiner, Notropis heterolepis…………………………………..…..291

Spottail Shiner, Notropis hudsonius………………………………………….296

Carmine Shiner, Notropis percobromus…………………………………..….301

Silverband Shiner, Notropis shumardi…………………………………..……306

Sand Shiner, Notropis stramineus……………………………………..……..310

Topeka Shiner, Notropis topeka……………………………………………...314

Channel Shiner, Notropis wickliffi…………………………………..………..320

Suckermouth Minnow, Phenacobius mirabilis……………………………….323

Bluntnose Minnow, Pimephales notatus……………………………………..327

Fathead Minnow, Pimephales promelas…………………………………...…332

Flathead Chub, Platygobio gracilis…………………………………………..338 xii

Western Blacknose Dace, Rhinichthys obtusus……………………..………..345

Longnose Dace, Rhinichthys cataractae……………………………………...350

Rudd, Scardinius erythrophthalmus………………………………...………..356

Creek Chub, Semotilus atromaculatus………………………………….…….361

CHAPTER 11: FAMILY CATOSTOMIDAE………………………………….……367

River Carpsucker, Carpiodes carpio……………………………...………….370

Quillback, Carpoides cyprinus……………………………….………………375

Highfin Carpsucker, Carpiodes velifer………………………………….……380

Longnose Sucker, Catostomus Catostomus………………………….……….385

White Sucker, Catostomus commersonii……………………………………..390

Mountain Sucker, Pantosteus jordani……………………………….………..394

Blue Sucker, Cycleptus elongatus……………………………………...……..399

Smallmouth Buffalo, Ictiobus bubalus……………………………...………..403

Bigmouth Buffalo, Ictiobus cyprinellus………………………………..……..408

Black Buffalo, Ictiobus niger…………………………………………..……..413

Silver Redhorse, Moxostoma anisurum…………………………………...….417

River Redhorse, Moxostoma carinatum…………………………………..….421

Golden Redhorse, Moxostoma erythrurum………………………………..….426 xiii

Shorthead Redhorse, Moxostoma macrolepidotum……………………….….431

Greater Redhorse, Moxostoma valenciennesi……………………………..….436

CHAPTER 12: FAMILY ICTALURIDAE…………………………………….…….441

Black Bullhead, Ameiurus melas………………………………………….….443

Yellow Bullhead, Ameiurus natalis…………………………………………..447

Brown Bullhead, Ameriurus nebulosus………………………………………451

Blue Catfish, Ictalurus furcatus……………………………………..………..455

Channel Catfish, Ictalurus punctatus…………………………………………459

Stonecat, Noturus flavus…………………………………………………..….464

Tadpole Madtom, Noturus gyrinus……………………………………..…….470

Flathead Catfish, Pylodictis olivaris………………………………………….475

CHAPTER 13: FAMILY ESOCIDAE……………………………………………….479

Grass Pickerel, Esox americanus……………………………………………..481

Northern Pike, Esox Lucius………………………………………………...…486

Muskellunge, Esox masquinongy………………………………………….….491

Central Mudminnow, Umbra limi………………………………………….…495

CHAPTER 14: FAMILY SALMONIDAE……………………………………….….499

Cisco, Coregonus artedi………………………………………………...……502 xiv

Lake Whitefish, Coregonus clupeaformis………………………………...….508

Cutthroat Trout, Oncorhynchus clarkia……………………………..………..514

Rainbow Trout, Oncorhynchus mykiss…………………………………...…..518

Chinook Salmon, Oncorhynchus tshawytscha…………………………….….523

Brown Trout, Samo trutta…………………………………………………….528

Brook Trout, Salvelinus fontinalis……………………………………………535

Lake Trout, Salvelinus namaycush…………………………………………...541

CHAPTER 15: FAMILY OSMERIDAE…………………………………………….545

Rainbow Smelt, Osmerus mordax…………………………………………....547

CHAPTER 16: FAMILY PERCOPSIDAE…………………………………………..553

Trout-perch, Percopsis omiscomaysus……………………………………..…555

CHAPTER 17: FAMILY GADIDAE……………………………………………...…559

Burbot, Lota lota…………………………………………………….………..561

CHAPTER 18: FAMILY FUNDULIDAE…………………………………..…….…566

Banded Killifish, Fundulus diaphanous…………………………….………..568

Plains Topminnow, Fundulus sciadicus………………………………...……572

Northern Plains Killifish, Fundulus zebrinus……………………………..….577

CHAPTER 19: FAMILY GASTEROSTEIDAE………………………………..……582 xv

Brook Stickleback, Culaea inconstans………………………………...……..584

CHAPTER 20: FAMILY MORONIDAE……………………………………………588

White Bass, Morone chrysops………………………………………………..590

CHAPTER 21: FAMILY CENTRARCHIDAE…………………………………..….595

Rock Bass, Ambloplites rupestris………………………………………….…598

Green Sunfish, Lepomis cyanellus……………………………………………603

Pumpkinseed, Lepomis gibbosus……………………………………………..608

Orangespotted Sunfish, Lepomis humilis…………………………………..…613

Bluegill, Lepomis macrochirus…………………………………………….…617

Redear Sunfish, Lepomis microlophus……………………………………….623

Smallmouth Bass, Micropterus dolomieu…………………………………….628

Largemouth Bass, Micropterus salmoides……………………………………633

White Crappie, Pomoxis annularis………………………………………...…638

Black Crappie, Pomoxis nigromaculatis………………………………..…….643

CHAPTER 22: FAMILY PERCIDAE…………………………………………...…..647

Iowa Darter, Etheostoma exile………………………………………………..649

Johnny Darter, Etheostoma nigrum…………………………………….…….653

Yellow Perch, Perca flavescens……………………………………………....658 xvi

Logperch, Percina caprodes………………………………………………….666

Blackside Darter, Percina maculate…………………………………...……..671

Slenderhead Darter, Percina phoxocephala………………………………….676

River Darter, Percina shumardi………………………………………..……..680

Sauger, Sander canadensis………………………………………………...…685

Zander, Sander lucioperca……………………………………………..……..691

Walleye, Sander vitreus………………………………………….………...…695

CHAPTER 23: FAMILY SCIAENIDAE……………………………………...……..702

Freshwater Drum, Aplodinotus grunniens……………………………..……..704

CHAPTER 24: FAMILY CICHLIDAE………………………………………..…….709

Jack Dempsey, Rocio octofasciata………………………………………...…711

xvii

LIST OF FIGURES

Figure 1. Map illustrating where the Northern Divide separates a large portion of the

Dakotas into two main watersheds: the Hudson Bay and the Gulf of Mexico. The

Dakotas are also made up of eight Environmental Protection Agency (EPA) Level II

Ecoregions (Bryce et al. 1996). (Map courtesy of Nick

Kludt)…………………………….……………………………………...………………3

Figure 2. Map illustrating the major river basins in North Dakota and South Dakota.

(Map courtesy of Nick

Kludt)………………………………………………………………………...………...17

Figure 3. Map illustrating the six Missorui River Reservior dams constructed by the

U.S. Army Corps of Engineers……………………………………………………..….25

xviii

ABSTRACT

FISHES OF THE DAKOTAS

KATHRYN E. SCHLAFKE

2020

Biological reference materials play a critical role in studies of species biodiversity, biology, distribution, and historic diversity. Since the publications Fishes of

North Dakota (Hankinson, 1929) and Fishes of South Dakota (Bailey and Allum, 1962), no new books have attempted to update the knowledge gained from the past decades of fish surveys and incorporate georeferenced collections at the state level. The last published book with the most complete accounts of fishes from North Dakota and South

Dakota is the Distribution of Fishes in North and South Dakota Basins affected by the

Garrison Diversion Unit (Owen et al. 1981) which includes point distribution maps, short species descriptions, and a dichotomous key of 105 species. Since these publications, several species have been introduced or discovered in one or both states. This project aims to create a new reference for all species of fish in North Dakota and South Dakota by synthesizing data from both states. The Fishes of the Dakotas book will include detailed point distribution maps, an updated taxonomic key, color plates or illustrations, and complete species accounts for 125 fishes. Species accounts, or literature reviews, in this new biological reference will include etymology, morphology, distribution and habitat, reproduction, food and feeding, and age and growth. This reference will be a valuable tool for students, researchers, biologists, and enthusiasts.

xix

This thesis was written in preparation for the forthcoming publication of the Fishes of

the Dakotas book, that will include additional chapters on how to identify fishes, a

dichotomous key, species color photographs, as well as point distribution maps.

1

CHAPTER 1

WATERS AND GEOLOGY OF NORTH DAKOTA AND SOUTH DAKOTA

North Dakota and South Dakota, often referred to together as the “Dakotas”, lie entirely within the Great Plains region of North America and have the renowned Missouri

River meandering through from the northwest to the southeast. The Missouri River once long served as a boundary line for several territories during the Westward Expansion. Not yet declared as two individual states, the land which would eventually become North

Dakota and South Dakota was included in part of the Louisiana Territory, which was purchased from the United States from France in 1803 by the Louisiana Purchase. From

1812 to 1861, the section of land west of the Missouri River was then included in the

Nebraska Territory, and the section of land east of the Missouri River belonged successively to the territory of Missouri (1812-1834), Michigan (1824-1836), Wisconsin

(1836-1838), Iowa (1838-1849) and Minnesota (1849-1858) (Visher, 1918). Between

1858 and 1861, the area of land was considered unorganized and became known as

“Land of the Dakotas” (Visher, 1918). In 1861, the Dakota Territory was then established to include what are now North Dakota, South Dakota, the majority of Montana, and portions of Nebraska and Wyoming (Visher, 1918) combined in 1863, until 1889 when the territory was split and admitted to the Union as two separate states: North Dakota and

South Dakota.

To many, North Dakota and South Dakota are recognized as vast, flat, wide-open regions of prairie with gently rolling hills that seem nearly endless when glaring out ones windshield driving across either state. However, if ventured off the beaten path, or in this case the interstate, one would realize that North Dakota and South Dakota are also 2 characterized by having deep, broad or narrow valleys above large rivers, natural lakes and large impoundments tucked behind hillsides or bluffs, and several streams winding through remote areas of prairie, desert buttes, or forested areas. Like many other regions within the United States, the explanation for the creation of these different geological features that we see today are due to past glacial movements and patterns.

One geological feature that often goes unnoticed by many is the Northern Divide, or commonly referred to as the Laurentian Divide in northern Minnesota (Gonzalez,

2003). Many people are under the misconception that there is only one continental divide in the United States and North America that occurs along the crest of the Rocky

Mountains (Gonzalez, 2003). This continental divide, often incorrectly referred to as the

Continental Divide and correctly referred to as the Great Divide, is only one of four distinct continental divides that occur within the United States and North America. The three other continental divides are the Northern Divide, the Eastern Divide, and the St.

Lawrence Seaway Divide.

The Northern Divide originates deep within the interior of the North American continent and shares the same path as the Great Divide from Seward Peninsula to Triple

Divide Peak, Montana. From Triple Divide Peak, the Northern Divide extends east through Montana, Alberta, Saskatchewan, North Dakota, South Dakota, and northern

Minnesota until it continues north and east across Ontario and Quebec (Gonzalez, 2003).

The Northern Divide separates a large portion of North America, and specifically the

Dakotas, into two main watersheds: the Hudson Bay and the Gulf of Mexico (Figure 1).

The Hudson Bay and Gulf of Mexico watersheds drain into different oceans or seas on different sides of the continent. 3

Figure 1: The Northern Divide separates a large portion of the Dakotas into two main watersheds: the Hudson Bay and the Gulf of Mexico. The Dakotas are also made up of eight

Environmental Protection Agency (EPA) Level II Ecoregions (Bryce et al. 1996).

4

The Hudson Bay watershed primarily encompasses the central territories of

Canada, but also extends south into Montana, North Dakota, South Dakota, and northern

Minnesota. In North Dakota, rivers and streams that occur within the Souris, Sheyenne, and Red River of the North drainages flow north, eventually leading to the Hudson Bay.

In South Dakota, the Hudson Bay watershed extends into the extreme northeastern corner and streams and rivers within this region called the Bois de Sioux River drainage which includes the Red River of the North drainage in South Dakota also flow north to the

Hudson Bay (Figure 1).

The Gulf of Mexico watershed covers a vast majority of the United States and comprises the Mississippi River watershed. The Mississippi River drainage roughly spans over the southwestern half of North Dakota and the majority of South Dakota (>99%)

(Hoagstrom et al., 2007). The Mississippi River drainage in the Dakotas is divided into two major sub-drainages: the Upper Minnesota River drainage and the Upper Missouri

River drainage. The Minnesota River drainage covers less than 3% of South Dakota and is drained by the Upper Minnesota River (Hoagstrom et al., 2007). The Upper Missouri

River drainage spans across the majority of central and eastern Montana, the northern half of Wyoming, the southwestern half of North Dakota, approximately 97% of South

Dakota, the northern border of Nebraska, and the northwestern corner of Iowa (Goolsby et al., 1999; Hoagstrom et al., 2007).

In North Dakota, rivers and streams to the west, southwest, and south-central region of the Northern Divide such as the James, Cannonball, Heart, Knife, and Little

Missouri Rivers, flow into the Missouri River, which conjoins with the Mississippi River eventually flowing into the Gulf of Mexico. All the rivers and streams within South 5

Dakota besides those that occur within the Red River of the North and the Minnesota

River drainages also flow into the Missouri and eventually empty into the Gulf of Mexico via the Mississippi River (Figure 1). The Devils Lake basin in North Dakota is recognized today as a closed basin, since there is currently no external drainage that leads to either the Hudson Bay or Gulf of Mexico watersheds (Gonzalez, 2003). The basin is only temporarily closed off to either watershed, as water may spill over into an externally draining river if water levels rise high enough (Gonzalez, 2003).

Despite the differences in the direction of flow between the Hudson Bay and the

Gulf of Mexico watersheds, the fish assemblages between the two are remarkably similar.

This is large in part due to the historical geology of the region. Four main time periods, or eras (Precambrian, Paleozoic, Mesozoic, and Cenozoic), help break apart our earth’s history. The most recent era, the Cenozoic, is split into two main periods, the Tertiary and the Quaternary. The Tertiary period began about 65 million years ago and lasted until roughly 1.8 million years ago when the Quaternary period began, which is the era we continue to live in today. Most of the later Quaternary period is often referred to as the

“Ice Age” or Pleistocene Epoch which took place roughly 2,588,000-11,700 years ago, just prior to the Holocene Epoch, which we currently live in today (Hendon & Matthews,

2014). During the Pleistocene Epoch, the formation of glaciers or large sheets of ice occurred, spanning from northern Canada to the northern and central portions of the

United States. The glacier or sheet of ice that covered the majority of North Dakota, and the eastern portion of South Dakota was called the Wisconsinan glacier.

The Wisconsinan glacier formed roughly 40,000 years ago, and advanced and retreated several times until it melted entirely roughly 12,000 years ago (Herman & 6

Johnson, 2012). As the glacier advanced, fish species were driven into the southern parts of the Mississippi River drainage, and as the glaciers retreated, the fish species were able to recolonize within the waterways along the north and south corridors (Oberdorff et al.,

1995). More specifically, fish species indigenous to the ancient and unaltered Missouri

River and its main tributaries that once belonged to the Hudson Bay drainage, were now free to roam within the Mississippi River drainage, and vice versa (Owen et al., 1981).

The historical back-and-forth shifting patterns of the glaciers during the

Pleistocene Epoch have contributed to today’s trend of having lower fish species diversity, or richness, in northern and western states, versus a greater diversity of fishes in states to the south and east. An example of this trend would be the Missouri River.

Native species richness increases with downstream movement, with 64 species in

Montana to 110 species in Missouri (Galat et al., 2005). This statement also applies to the

Dakotas, specifically with lower species diversity on the western half of the states compared to the eastern half (Hoagstrom et al., 2007). These spatiotemporal changes of fishes reflect interactions between natural factors such as climate, physiography, hydrology, and zoology, as well as anthropogenic factors like impoundment, flow and temperature alterations, and introduced species (Galat et al., 2005). Although fish assemblages within the Hudson Bay and Gulf of Mexico watersheds are rather similar, the distribution and abundance of fishes within the Dakotas are continually changing due to species range expansions and species declines (Hoagstrom et al., 2007).

Before we dive any further into the fishes aspect of the book, it is important for one to know more specifics about the ecoregions and drainages that occur throughout the

Dakotas. These regions help define the waterways within both states, and often help 7 explain why some species of fish occur where they do. As mentioned previously, the period of glaciation during the Pleistocene Epoch helped form and construct the current landscape of North America and specifically, the Dakotas, as we know it today.

Level III Ecoregions

Currently, the Dakotas are broken up into eight different Level III Ecoregions by the Environmental Protection Agency (EPA) that help to better categorize areas with similarities in the type, quality, and quantity of resources in both states (Bryce et al.,

1996). Out of the 98 total Level III Ecoregions that span across the entire United States, eight occur within North Dakota and South Dakota (Bryce et al., 1996) (Figure 1). The eight regions that span across the Dakotas east to west include: The Lake Agassiz Plain,

Western Corn Belt Plains, Northern Glaciated Plains, Northwestern Glaciated Plains,

Northwestern Great Plains, Nebraska Sand Hills, High Plains, and the Middle Rockies

(Figure 1).

Lake Agassiz Plain

After the Wisconsinan glacier had melted, glaciers directly to the north in Canada still existed and essentially formed a large dam that blocked any rivers trying to flow north (Herman & Johnson, 2012). Overtime, all the collected water that was not allowed through by the glaciers formed a massive lake called Lake Agassiz. Lake Agassiz was the largest of the many Pleistocene lakes in North America. The majority of Lake Agassiz was located in Canada in what is now Saskatchewan, Manitoba, and Ontario. Only a small fraction (about one-fifth) of the area of Lake Agassiz was located within the United

States in what is now North Dakota, South Dakota and Minnesota (Upham, 1895). 8

Lake Agassiz, named after the Swiss geologist Louis Agassiz, was roughly 700 miles long, 200 miles wide, and 300 feet deep (Herman & Johnson, 2007). During storms, large crashing waves helped define the shorelines which are now easily recognized and mapped across the prairie region. As the lake slowly retreated and diminished from south to north over time, it left behind an extremely flat portion of land with very fertile soil, which makes favorable conditions for agriculture. The very southwestern shoreline of Lake Agassiz now makes up the western border of the Red

River Valley in the Dakotas (Upham, 1895). The lake remnants of Lake Agassiz are lakes

Winnipeg, Manitoba, and Winnipegosis (Upham, 1895).

Level III Ecoregion 48, the Lake Agassiz Plain is located on the eastern edge of

North Dakota, and also extends into a small portion in South Dakota in the extreme northeast corner of the state (Figure 1). Today, the Lake Agassiz Plain is a low-relief area covering a total of 7,101 square miles and is made up of flat to low rolling plains covered by moraine and lacustrine deposits, with calcareous glacial till found in the northern region (Griffith, 2010). The thick beds of lacustrine sediments on top of glacial till have created an extremely flat landscape with relatively few lakes and wetlands. Once heavily comprised of tallgrass prairie and riparian areas, the land has now been extensively converted to row crop agriculture (Bryce et al., 1996; Griffith, 2010). The hydrology of the region is represented by low densities of low gradient stream and river networks that all drain into the Red River of the North (Griffith, 2010).

Western Corn Belt Plains

Level III Ecoregion 47, the Western Corn Belt Plains includes most of the grass- dominated land cover in the Corn Belt region that spans a total of 1,453 square miles over 9 the Midwest, and encompasses five Midwestern states: South Dakota, Minnesota, Iowa,

Nebraska and Kansas (Bryce et al., 1996). The ecoregion is located in the southeastern corner of South Dakota, and is characterized by a temperate climate and fertile soils of nearly level to gently rolling glaciated till plains that lie over Mesozoic and Paleozoic deposits of shale, sandstone, and limestone (Bryce et al., 1996; Griffith, 2010). Once vastly filled with tall grass prairie and small areas of oak woodlands, approximately three-fourths of the ecoregion has now been converted for intense row crop agriculture, with the remainder being pastureland including bits of native prairie and anthropogenically modified grasses and legumes (Griffith, 2010; Omernik & Gallant,

1988; Wright & Wimberly, 2013). A few natural lakes, and a system of intermittent and perennial streams that typically have shallow, narrow valleys due to channelization from agricultural practices characterize the ecoregions hydrology (Griffith, 2010). Patches of remaining deciduous woodlands also sometimes border these prairie streams (Omernick

& Gallant, 1988). In South Dakota, the ecoregion contains parts of the lower Big Sioux

River, lower Vermillion River, lower James River and the Missouri River. The Western

Corn Belt Plains also converges with much of the Prairie Pothole Region (PPR), an area that provides the most productive breeding habitat for migrating waterfowl in North

America (Batt et al. 1989, Wright & Wimberly, 2013).

Northern Glaciated Plains

Level III Ecoregion 46, the Northern Glaciated Plains span from Saskatchewan and Manitoba, Canada in the north, and south to north-central North Dakota, eastern

South Dakota and southwest Minnesota (Omernik & Gallant, 1988). The terrain of the sub-humid, 48,511 square miles ecoregion is characterized by nearly level to gently 10 rolling glacial till plains, and areas of glacial lacustrine and ridged fluvial deposits, over

Tertiary and Cretaceous sandstone and shale (Bryce et al., 1998; Griffith, 2010).

Historically covered with mixtures of short and tallgrass prairie, and trembling aspen and oak groves, much of the landscape has now been converted to farmland (cattle grazing, livestock production, and irrigated agriculture) given the ecoregions very fertile and productive soil (Griffith, 2010; Omernick & Gallant, 1988). Areas of native woodlands, pastures, and seasonal wetlands also occur in the area, providing multiple habitat uses to wildlife (Bryce et al., 1988; Griffith, 2010). Low densities of intermittent streams stemming from larger rivers, high concentrations of temporary and seasonal prairie pothole wetlands, and many small glacial or kettle lakes scattered across the landscape make up the ecoregion’s hydrology (Griffith, 2010; Omernick & Gallant, 1988). More specifically, the intermittent waterways and wetlands that occur throughout the grassland landscape drain to the prairie pothole basins (Bryce et al., 1998; Krause et al., 2013). In the Dakotas region, the Northern Glaciated Plains contain the headwaters of numerous tributaries to the Red River of the North, as well as large portions of the Souris,

Sheyenne, James, Vermillion, Big Sioux, and Minnesota Rivers.

Northwestern Glaciated Plains

Level III Ecoregion 42, the Northwestern Glaciated Plains, is a 28,378 square mile ecoregion that spans from Saskatoon in the north, to southwestern Saskatchewan, southeastern Alberta, and south to northern Montana, the central region of the Dakotas, and into the northern portion of Nebraska (Griffith, 2010). In North Dakota and South

Dakota, the Northwestern Glaciated Plains lie entirely east of the Missouri River, and represent the westernmost extent of continental glaciation during the Pleistocene Epoch. 11

The ecoregion is essentially a transitional region between the more level, moist, and agricultural Northern Glaciated Plains to the east, and the dryer and more irregular

Northwestern Great Plains to the west and southwest (Bryce et al., 1996; Chapman et al.

2001; Griffith, 2010).

The region is characterized by nearly level plains and gently rolling hills that were once covered with native tall grasses and shrubs, over shale, sandstone, and limestone soils that were derived from glacial till left by the Wisconsinan glacier (Bryce et al.,

1996; Griffith, 2010). The hydrology of the area consists of high concentrations of intermittent streams, with some perennial streams and larger rivers present on the western edge of the ecoregion (Griffith, 2010). These waterways are drained by the Missouri

River system to the south and the Saskatchewan River to the north (Griffith, 2010). In some portions of the ecoregion, the landscape is scattered with high concentrations of seasonal and semi-permanent wetlands, often referred to as Prairie Potholes, which are heavily utilized by waterfowl for nesting and migration purposes creating prime opportunities for waterfowl hunting conditions (Griffith, 2010). Today, much of the land is utilized as dryland agriculture such as cropland and rangeland for cattle grazing (Bryce et al. 1996; Griffith, 2010).

Northwestern Great Plains

Level III Ecoregion 43, the Northwestern Great Plains, is an entirely unglaciated region of land that extends from the base of the Rocky Mountains in Montana, east to the southwestern corner of North Dakota, most of western South Dakota (excluding the

Middle Rockies, the High Plains, and the Nebraska Sandhills), and into the northeast corner of Wyoming. In the Dakotas, the ecoregion lies entirely west of the Missouri 12

River, and is bordered by the Missouri River and Northwestern Glaciated Plains in the east, and surrounds the Nebraska Sand Hills, the High Plains, and the Middle Rockies in southwestern South Dakota.

The ecoregion is largely characterized by gently rolling plains and occasional buttes, as well as some dissected areas of badland terrain and steep river breaks over

Cretaceous shale and Tertiary sandstone deposits (Bryce et al., 1996; Griffith, 2010).

Mixed-grass prairie and shrub land are the primary types of vegetation that cover the area within Montana and the Dakotas, with more shortgrass prairie occurring throughout

Wyoming (Biondini et al., 1998; Tan et al., 2005). The once vast native shortgrass and mixed grass prairies in the ecoregion now largely persist as rangeland areas used for livestock grazing, as well as some areas of cropland on more level grounds (Bryce et al.

1996; Griffith, 2010). The hydrology of the ecoregion is characterized by mostly intermittent and ephemeral streams, with a few perennial rivers that drain west into the

Missouri River (Griffith, 2010). The western tributaries of the Missouri River within the ecoregion spanning throughout the Dakotas are the lower Yellowstone, Little Missouri,

Knife, Heart, Cannonball, Grand, Moreau, Cheyenne, Bad, White, upper Niobrara, and the Keya Paha rivers. The majority of these rivers have uniquely turbid waters that are seldom clear due to the underlying deposits of sediments and clays the waters carry from erosion and weathering rock upstream near the Badlands (Bryce et al. 1996). Small impoundments also occur semi-frequently over the landscape, as well as some large reservoirs along the Missouri River (Griffith, 2010).

13

Nebraska Sand Hills

Level III Ecoregion 44, the Nebraska Sandhills, is one of the most distinct and homogenous ecoregions in North America occurring within the heart of the Great Plains region, and spans over north-central and northwestern Nebraska, as well as a small region along the south-central edge of South Dakota (Griffith, 2010). In South Dakota, the ecoregion is bordered by the Northwestern Great Plains and falls west of the Missouri

River. The Nebraska Sand Hills were formed during periods when climactic changes killed the native grasses and revealed underlying sand to become exposed (Billesbach &

Arkebauer, 2012).

The ecoregion is now recognized as one of the largest areas of grass stabilized sand dunes in the world, and lies on top of the deepest part of the sand and gravel deposits that make up the High Plains or Ogallala Aquifer, which supplies roughly 30% of all the commercial agricultural irrigation water used annually in the United States

(Bryce et al, 1996; USGS, 1997; Bleed & Flowerday, 1998; Chapman et al., 2001). The vastly barren terrain lies on rolling to steep, irregular, wind-sculpted sand dunes over

Tertiary sandstones and conglomerates that lie beneath Quaternary aeolian sand (Griffith,

2010). The vegetation within the vastly treeless ecoregion mainly occurs within the valleys of the sand dunes, and is primarily grass stabilized with mixtures of short, mid, and tallgrass prairie, as well as some riparian areas in the north and east where some trees are present (Billesbach & Arkebauer, 2012). Numerous lakes, alkaline wetlands, and a general lack of streams make up the Nebraska Sand Hills hydrology (Griffith, 2010). A few larger perennial streams and small rivers such the Niobrara River also flow within the region. Due to the soil composition, row crop agriculture is scarce across the 14 landscape, and much of the land is instead used for ranching and livestock grazing

(Griffith, 2010).

High Plains

Level III Ecoregion 25, the High Pains, covers only a small area near the south- central edge of South Dakota, but spans a rather large latitudinal extent from southeastern

Wyoming, western Nebraska and eastern Colorado, south to western Kansas, Oklahoma and Texas, and into eastern New Mexico (Griffith, 2010). In South Dakota, the ecoregion is bordered by the Northwestern Great Plains and falls west of the Missouri River. The

High Plains were originally formed by the erosion of the Rocky Mountains leaving behind nearly smooth to slightly irregular rolling plains and tablelands that lie over

Tertiary and Cretaceous sandstones, siltstones, claystones, and caliche layers (Bryce et al., 1996; Griffith, 2010). Vegetation across the landscape is primarily comprised of drought resistant short and mid mixed grass prairies, and the small part of the ecoregion that intersects into South Dakota contains ponderosa pine (Bryce et al., 1996).

Intermittent and ephemeral streams in the northern and central portions of the ecoregion make up the majority of the hydrology characteristics of the High Plains, as much of the southern portion of the ecoregion has little to no streams (Griffith, 2010). Some larger rivers like the Platte River also occur within the ecoregion. Today, the landscape is used for agricultural purposes such as grazing and cropland. Most of the farming practices that occur within the area utilize water that is irrigated by the Ogallala Aquifer since moisture is limited in the High Plains ecoregion (Bryce et al., 1996).

15

Middle Rockies

Level III Ecoregion 17, the Middle Rockies, spans over most of southwestern

Montana, eastern Idaho, and northern Wyoming; however, the ecoregion also includes an outlier, or “island” portion of land that shares the same characteristics in northeastern

Wyoming and western South Dakota known as the Black Hills (Bryce et al., 1996;

Griffith, 2010). The Middle Rockies ecoregion is geologically characterized as a dome- shaped uplift made up of a mixture of geological features such as high elevations of alpine mountain ranges, grassy glacial basins, and plateaus (DeWitt et al. 1989; Bryce et al. 1996). These geological features all lie over a variety of rock types and ages spanning throughout the ecoregion.

The Middle Rockies were created roughly 60-65 million years ago, with the center of the dome comprised of Precambrian metamorphic rock surrounded by

Cretaceous sedimentary formations that completely encircle the core (DeWitt et al. 1989;

Williamson & Carter, 2001). Vegetation within the ecoregion, specifically within the area that occurs in South Dakota, consists of ponderosa pine forests, partly wooded and shrub- covered foothills, alpine grasslands, and meadows (Griffith, 2010). A mixture of small alpine glacial lakes, larger lakes, and high gradient perennial streams make up the varying hydrological features of the Middle Rockies (Griffith, 2010). The sedimentary formations and underlying geology of the ecoregion have a great effect on the area’s hydrology (Williamson & Carter, 2001). In the Black Hills, limestone formations that lie west of the center act as groundwater recharge zones with little amounts of discharge

(Williamson & Carter, 2001). The surface discharge of streams that originate in the center of the Black Hills is primarily lost as they flow north or east away from the core 16

(Williamson & Carter, 2001). Streams within the region have variable morphology and biological characteristics, but generally have cobble and rubble substrates, with slack water portions capable of accumulating fine sediments and organic debris (Schultz,

2011). Water quality in numerous Black Hills streams have been historically impacted by mining activities within the last century (Rahn et al., 1996). Land use in the ecoregion includes forestry, ranching, grazing, recreation, and mining.

Major River Basins of North Dakota and South Dakota

Currently, North Dakota and South Dakota are broken up into 20 different basins within the eight EPA Level III Ecoregions (Figure 2). Just like the ecoregions, each basin is a sub-drainage, and belongs to either the Hudson Bay or Gulf of Mexico watersheds in which they eventually drain to. This next section will introduce each of the 20 basins (in alphabetical order) and will describe the main hydrological features of each basin in more detail. 17

Figure 2: Major River basins in North Dakota and South Dakota.

18

Bad River Basin (Missouri River Sub-drainage)

The Bad River basin located in west-central South Dakota drains roughly 8,622 km2 (3,176 mi2) of land (Figure 2). The basin is surrounded by the Missouri River basin in the northeast, the Cheyenne River basin in the northwest, and the White River basin in the south. Eventually draining into the Missouri River, the upper portion, or North Fork of the Bad River gets its water from the Badlands and artesian wells near Philip, South

Dakota, where it meets up with the South Fork. The South Fork of the Bad River is sourced from Big Buffalo Creek and Whitewater Creek in the White River Badlands

(Hoagstrom et al., 2007). These sources contribute minimal flow, which contributes to the rivers main characteristic as having little to no constant river flow (Pirner, 2018).

From the convergence of the North and South Forks, the mainstem of the Bad River then flows east, where it eventually converges with the Missouri River near Fort Pierre, South

Dakota (Hoagstrom et al., 2007). Major tributaries of the Bad River basin include Dry

Creek, Plumb Creek and White Willow Creek (Hoagstrom et al., 2007).

Belle Fourche River Basin (Missouri River Sub-drainage)

The Belle Fourche River basin located in western South Dakota between the

Cheyenne and Moreau River basins drains roughly 8,509 km2 (3,285 mi2) of land (Figure

2). The upper portion of the basin was historically used for several hard-rock mining operations, one of which is still in production today (Pirner, 2018). The middle and lower parts of the river basin are now primarily used for irrigation and livestock watering. The

Belle Fourche River is the main river within the basin and inherits its name from the

French meaning “beautiful fork”. The headwaters of the Belle Fourche River are located in northeastern Wyoming, where it flows northeast until it crosses the Wyoming-South 19

Dakota border northwest of Belle Fourche, South Dakota and continues to flow southwest along the northern border of the Black Hills until it converges with the

Cheyenne River east of Boneita Springs, South Dakota.

Big Sioux River Basin (Missouri River Sub-drainage)

The Big Sioux River basin in eastern South Dakota drains roughly 10,056 km2

(26,044 mi2) of land and is the most populated river basin in the state (Figure 2)

(Hoagstrom et al., 2007; Pirner, 2018). The basin is surrounded by the Minnesota River basin in the northeast, the James River basin in the west, and the Vermillion River basin in the south. The lower portion of the Big Sioux River serves as the Iowa-South Dakota boarder, where it eventually converges with the Missouri River in Sioux City, Iowa.

Cannonball River Basin (Missouri River Sub-drainage)

The Cannonball River basin located west of the Missouri River in southwest central North Dakota, and northwest central South Dakota drains roughly 11,184 km2

(4,318 mi2) of land (Figure 2). The basin is surrounded by the Missouri River basin in the east, the Heart River in the north, the Little Missouri River basin in the west, and the

Grand River basin in the south. The main river within the basin, the Cannonball River, originates from White Lake, just east of Amidon, North Dakota. From White Lake, the

Cannonball River flows northeast and then southeast until it drains into the Missouri

River near Cannonball, North Dakota.

Cheyenne River Basin (Missouri River Sub-drainage)

The Cheyenne River basin located west of the Missouri River in southwestern

South Dakota drains roughly 38,455 km2 (14,847 mi2) of land in South Dakota (Figure 2)

(Hoagstrom et al., 2007). The entire Cheyenne River basin spans approximately 56,047 20 km2 (21,639 mi2) within Wyoming, Montana, South Dakota, and Nebraska (Duehr, 2004;

Galat et al., 2005). Within South Dakota, the basin extends through diverse landscapes including the Black Hills and the Badlands, and is surrounded by the Belle Fourche,

Moreau, Bad, and White River basins. The water quality within the basin is generally poor, due to both natural and agricultural practices and highly erodible soils contribute to high levels of sedimentation during heavy rainfall (Pirner, 2018). The Cheyenne River is made from two main arms (Upper Cheyenne and Belle Fourche) that flow to the north and south around the Black Hills and converge on the northeastern side of the Black

Hills. After the convergence point that occurs just west of Pedro, South Dakota, the lower portion of the Cheyenne River flows east roughly 386 km (240 mi) before draining into

Lake Oahe (Duehr, 2004). The Upper Cheyenne River also flows through Angostura

Reservoir near Cascade Springs, South Dakota. A unique feature of the Cheyenne River basin is the Fall River, as it is fed by hot springs.

Devils Lake River Basin

The Devils Lake basin located east of the Missouri River in northeast central

North Dakota drains roughly 9,938 km2 (3,837 mi2) of land (Figure 2). The basin is surrounded by the Red River of the North to the east and north, the Souris River basin to the west, and the Sheyenne River basin in the south. Once entirely glaciated, the receding ice formed what is now Devils Lake. The basin is named after Devils Lake, North

Dakota’s largest natural body of water, and the state’s second largest body of water after

Lake Sakakawea. Devils Lake, located in the southern end of the basin, is an endorheic, or closed lake, and naturally fluctuates from overflowing to dry. During overflow 21 periods, it naturally spills into Stump Lake, where the combined flows seep into the

Sheyenne River.

Grand River Basin (Missouri River Sub-drainage)

The Grand River basin located west of the Missouri River in southwest North

Dakota and in the central northwest region of South Dakota drains roughly 13,967 km2

(5,393 mi2) of land (Figure 2). The basin is surrounded by the Cannonball River basin in the north, the Little Missouri River basin in the west, the Moreau River basin in the south, and the Missouri River basin in the east. Within North Dakota, the basin drains approximately 2,395 km2 (925 mi2). Within South Dakota, the basin drains roughly

11,572 km2 (4,468 mi2). The North Fork of the Grand River begins south of the town

Rhame, in southwestern North Dakota. The North Fork flows east near the North Dakota-

South Dakota border where it flows through Bowman-Haley Lake, just west of Haley,

North Dakota, and further southeast into Shadehill Reservoir near Shadehill, South

Dakota. The South Fork of the Grand River, and the Little Grand River converge nwest of Buffalo, South Dakota, where the South Fork continues to flow northeast into

Shadehill Reservoir. The North and South Forks of the Grand River exit Shadehill on the east side of the lake, and converge to create the mainstem of the Grand River, which flows east until it drains into Lake Oahe near Little Eagle, South Dakota.

Heart River Basin (Missouri River Sub-drainage)

The Heart River basin located west of the Missouri River in the central southwest region of North Dakota drains roughly 8,676 km2 (3,350 mi2) of land (Figure 2). The basin is surrounded by the Missouri River basin in the east, the Knife River basin in the north, the Little Missouri River basin in the west, and the Cannonball River basin in the 22 south. It is suggested that the eastern half of the basin was once glaciated plateau land, given the number of hills that contain moraine remnants, and the western half of the basin was unglaciated plateau which has rolling plains broken up by buttes (Maderak, 1966).

James River Basin (Missouri River Sub-drainage)

The James River basin located east of the Missouri River in both east-central

North Dakota and South Dakota drains roughly 54,390 km2 (21,000 mi2) of land total, with roughly 21,755 km2 (8,400 mi2) in North Dakota, and roughly 32,633 km2 (12,600 mi2) in eastern South Dakota (Figure 2) (Owen et al., 1981; Shearer and Berry, 2002).

The basin is surrounded by the Vermillion, Big Sioux, Minnesota, Red River of the

North, and Sheyenne River basins in the east, and the Missouri River basin in the west.

Once entirely glaciated and formed from the Lake Dakota Plain from the Pleistocene glaciation period, the main river within the basin, the James River, is now one of the lowest gradient rivers in the United States, and is thus prone to flooding (Benson, 1983;

Shearer and Berry, 2002). The James River begins in central North Dakota, near the town of Fessenden, and extends roughly 1,202 km (747 mi) south until if converges with the

Missouri River near Yankton, South Dakota (Benson, 1983).

Knife River Basin (Missouri River Sub-drainage)

The Knife River basin located west of the Missouri River in west-central North

Dakota drains roughly 6,488 km2 (2,505 mi2) of land (Figure 2). The basin is surrounded by the Missouri River basin in the east and north, the Little Missouri River basin in the west, and the Heart River basin in the south. The main river within the basin, the Knife

River, stems from the Knife River and the Little Knife River, before it converges just west of Manning, North Dakota. The mainstem of the Knife River flows east until it 23 converges with the Missouri River near Knife, North Dakota, just south of Lake

Sakakawea. A main tributary of the Knife River, the Branch Knife River, extends from the mainstem just east of Marshall, North Dakota, and flows south until it reaches just west of Hebron, North Dakota.

Little Missouri River Basin (Missouri River Sub-drainage)

The Little Missouri River basin located west of the Missouri River in southwestern North Dakota and northwestern South Dakota drains roughly 13,940 km2

(5,382 mi2) of land (Figure 2). Within the Dakotas, the basin is surrounded by the

Missouri River basin to the north, the Knife, Heart, Cannonball, and Grand River basins to the east, and the Moreau and Belle Fourche River basins in the south. Within North

Dakota, the basin drains roughly 12,329 km2 (4,760 mi2), and roughly 1,611 km2 (622 mi2) in South Dakota.

Minnesota River Basin

The Minnesota River basin located east of the Missouri River in northeastern

South Dakota drains roughly 2,529 km2 (976 mi2) of land in South Dakota (Figure 2)

(Hoagstrom et al., 2007). The basin is surrounded by the Red River of the North basin in the north, the James and Big Sioux River basins to the west, and the Upper and Lower

Mississippi River basins to the east. The Minnesota River stems from Big Stone Lake on the Minnesota-South Dakota border and flows southeast through Minnesota to eventually converge with the Mississippi River at Fort Snelling, Minnesota. Main tributaries of the

Minnesota River basin in South Dakota include the Little Minnesota, Whetstone, Yellow

Bank, and Lac qui Parle Rivers.

24

Missouri River Basin (Missouri River Sub-drainage)

The Missouri River sub-basin, or valley, in the Dakotas runs from the northwest corner of North Dakota down the near center of the state, and essentially divides South

Dakota into almost equal eastern and western halves (Figure 2). The basin includes sections of land on both sides of the Missouri River, and drains roughly 87,612 km2

(33,827 mi2) of land in the Dakotas. The Missouri River basin in its entirety is the second largest river basin in the United States, (followed by the Mississippi River basin) draining about one sixth of the continental United States (Pegg et al. 2003). In the United States, the basin spans across 1,371,017 km2 (529,000 mi2), and roughly 25,122 km2 (9,700 mi2) in Canada (USACE, 1998; Galat et al., 2005).

The Missouri River basin of course surrounds the well-known Missouri River, however many people are unfamiliar with the fact that the Missouri River is the longest river in North America flowing 3,768 km (2,341 mi), followed by the Mississippi River which flows 3,544 km (2,202 mi) (Benke & Cushing, 2005; Galat et al., 2005). Often referred to as the “Big Muddy”, due to the rivers frequent flooding and shifting channels which create high levels of turbidity, the Missouri River originates near Three Forks

Montana at the confluence of the Madison, Jefferson, and Gallatin rivers (Galat et al.,

2005). From there, the Missouri River flows southeast across the midcontinent until its confluence with the Mississippi River in St. Louis, Missouri.

During the late 1930’s and the early 1940’s when the nation was undergoing a state of depression, six mainstem dams were constructed along the Missouri River

(Figure 3). These dams were constructed to control flooding, provide necessary depths for navigational purposes, generate power, irrigational purposes, and for water quality 25 management (Galat et al., 1996). Of these six dams, one occurs in northeast Montana

(Fort Peck Dam) and five occur along the stretch of the Missouri River that flows through

North Dakota and South Dakota (Figure 3).

Figure 3: Six Missouri Reservior dams constructed by the U.S. Army Corps of Engineers. Garrison Dam (Lake Sakakawea)

The Garrison dam, named after the town of Garrison, North Dakota,

located directly north of the dam, is the only impoundment of the Missouri River

located in North Dakota between the towns of Riverdale and Park City in the

central part of the state. Built by the U.S. Army Corps of Engineers, construction

of the dam began in 1946, and the closing of the dam occurred on April 15, 1953.

The dam has officially been operating since 1955. The Garrison dam spans a 26 length of 3,444 m (11,300 ft) (excluding the spillway), and impounds Lake

Sakakawea, the 3rd largest reservoir in the United States. Lake Sakakawea is 286 km (178 mi) long, with a shoreline length of 2,156 km (1,340) and a surface area of 154,590 ha (382,000 acres). Maximum depth of the reservoir is 55 m (180 ft)

(USACE, 2012).

Oahe Dam (Lake Oahe)

Oahe dam is located just north of Pierre, South Dakota in the central part of the state. Built by the U.S. Army Corps of Engineers, construction of the dam began in 1948, and the closing of the dam occurred on August 3, 1958. The dam has officially been operating since 1962. Oahe dam spans a length of 2,835 m

(9,300 ft) (excluding the spillway), and impounds Lake Oahe, the 4th largest reservoir in the United States. Lake Oahe is 372 km (231 mi) long, with a shoreline length of 3,621 km (2,250 mi), and a surface area of 83,406 ha (206,100 acres). Maximum depth of the reservoir is 62 m (205 ft) (USACE, 2012).

Big Bend Dam (Lake Sharpe)

Big Bend dam is located near Fort Thompson, South Dakota in the south- central part of the state, downstream of Lake Oahe. Built by the U.S. Army Corps of Engineers, construction of the dam began in 1959, and the closing of the dam occurred on July 24, 1963. The dam has officially been operating since 1964. Big

Bend dam spans a length of 3,222 m (10,570 ft) (excluding the spillway), and impounds Lake Sharpe, the 54th largest reservoir in the United States. Lake

Sharpe, named after the 17th South Dakota governor, Merrill Q. Sharpe, is 129 km

(80 mi) long, with a shoreline length of 322 (200 mi), and a surface area of 23,020 27 ha (56,884 acres). Maximum depth of the reservoir is 24 m (78 ft) (USACE,

2012).

Fort Randall Dam (Lake Francis Case)

Fort Randall dam is located near Pickstown, South Dakota in the southeastern part of the state, downstream from Big Bend dam. Built by the U.S.

Army Corps of Engineers, construction of the dam began in 1946, and the closing of the dam occurred on July 20, 1952. The dam has officially been operating since

1953. Fort Randall dam spans a length of 3,261 m (10,700 ft) (including the spillway), and impounds Lake Francis Case, the 11th largest reservoir in the

United States. Lake Francis Case, named after the former senator of South

Dakota, Francis Higbee Case, is 172 km (107 mi) long, with a shoreline length of

869 km (540 mi), and a surface area of 41, 278 ha (102,000 acres). Maximum depth of the reservoir is 43 m (140 ft) (USACE, 2012).

Gavins Point Dam (Lewis and Clark Lake)

Gavins Point Dam is located just west of Yankton, South Dakota in the southeastern part of the state, downstream from Fort Randall dam. Built by the

U.S. Army Corps of Engineers, construction of the dam began in 1952, and the closing of the dam occurred on July 31, 1955. The dam has officially been operating since 1955. Gavins Point dam spans a length of 2,652 m (8,700 ft) including the spillway), and impounds Lewis and Clark Lake, named after the explorers Merriweather Lewis and William Clark of the famous Lewis and Clark expedition. Lewis and Clark Lake, located on theborder of South Dakota and

Nebraska, is 40 km (25 mi) long, with a shoreline length of 144 km (90 mi), and a 28

surface area of 31,000 acres. It is the smallest of the Missouri River reservoirs.

Maximum depth of the reservoir is 14 m (45 ft) (USACE, 2012). Located just

upstream of Gavins Point Dam is the 63 km (39 mi) section of Missouri National

Recreational River that stretches westward to Fort Randall Dam, and just

downstream of the dam is the 95 km (59 mi) section of Missouri National

Recreational River the stretches east towards Ponca State Park in northeastern

Nebraska. These two stretches of the Missouri River are the only sections of the

entire river that remain undammed or unchannelized (least altered).

The sixth impoundment, or dam, along the Missouri River is the most northern of the six, and occurs in northeast Montana, impounding Fort Peck reservoir. Together, these six impoundments cover approximately half of the upper 2,500 km (1,553 mi) of the Missouri River (Morris et al., 1968). These human alterations such as the construction of impoundments and channelization, especially within the middle Missouri region that runs through the Dakotas, have significantly altered the daily natural flow regime over time (Pegg et al., 2003).

Moreau River Basin (Missouri River Sub-drainage)

The Moreau River basin located west of the Missouri River in northeastern South

Dakota drains roughly 8,105 km2 (3,129 mi2) of land (Figure 2) (Hoagstrom et al. 2007).

The basin is surrounded by the Grand River basin to the north, the Little Missouri and

Belle Fourche River basins to the west, and the Cheyenne River basin to the south. The mainstem of the Moreau River stems from the North Fork and the South Forks, which headwaters are located in the Short Pine Hills near the South Dakota-Montana border.

The North and South Forks converge to form the mainstem of the Moreau River near 29

Zeona, South Dakota, where it flows eastward until it converges with the Missouri River, south of Mobridge, South Dakota.

Niobrara River Basin (Missouri River Sub-drainage)

The Niobrara River basin expands from eastern Wyoming and runs along the unglaciated terrain of the northern border of Nebraska with a small portion extending into southcentral South Dakota. Within South Dakota, the Niobrara River basin drains roughly 3,218 km2 (1,242 mi2), and lies west of the Missouri River and is surrounded by the White River basin to the north, the Missouri River basin to the east, and the Loup and

Elkhorn River basins in Nebraska to the south (Figure 2) (Hoagstrom et al., 2007). The

Keya Paha River is the main tributary in South Dakota that flows southeast beginning near Hidden Timber, South Dakota, and eventually converges with the mainstem of the

Niobrara River west of Butte, Nebraska.

Red River of the North River Basin (Hudson Bay Watershed)

The Red River of the North basin located east of the Missouri River along the eastern border of North Dakota, and the extreme northeastern corner of South Dakota drains roughly 27,058 km2 (10,447 mi2) of land (Figure 2). The basin is surrounded by the Devils Lake, Sheyenne, and James River basins to the west, the Minnesota River basin in the south, and the Mississippi and Rainy River basins in Minnesota to the east.

Once entirely glaciated, the Red River of the North was formed by melting ice, which formed a deep channel within the broad and level valley of the basin (Horton and

Follansbee, 1906). The headwaters of the Red River stem from Lake Traverse which is located on the Minnesota-South Dakota border. The Red River of the North flows north 30 into Lake Winnipeg, near Manitoba, Canada where it eventually drains into the Hudson

Bay from the Nelson River.

Sheyenne River Basin (Hudson Bay Watershed)

The Sheyenne River basin located east of the Missouri River in east-central North

Dakota drains roughly 27,712 km2 (10,700 mi2) of land (Figure 2) (Owen et al., 1981).

The basin is surrounded by the Red River of the North basin to the east, Devils Lake and

Souris River basins to the north, and the Missouri and James River basins to the west.

The main river within the basin, the Sheyenne River, is approximately 885 km (550 mi) long and begins in the drift prairie, north of McClusky, North Dakota, and flows east until it converges with the North Fork of the Sheyenne River near East Fork Township,

North Dakota (Owen et al., 1981). The mainstem continues to flow east along the southern border of the Devils Lake basin, and then continues south until it flows into

Lake Ashtabula, behind Baldhill Dam, just north of Valley City, North Dakota. The river continues to flow until it turns northeast near Lisbon, North Dakota, and eventually converges with the Red River of the North, just north of Fargo, North Dakota.

Souris River Basin (Hudson Bay Watershed)

The Souris River basin located east of the Missouri River in north-central North

Dakota drains roughly 22,607 km2 (8,729 mi2) of land (Figure 2). The basin is surrounded by the Sheyenne and Devils Lake River basins in the east, the Red River of the North

River basin in the north, and the Missouri River basin in the southwest. The main river that flows through the basin, the Souris River (also known as the Mouse River), begins in southeast Saskatchewan, Canada, and flows south to Velva, North Dakota before it turns back north and flows into Manitoba, Canada, just north of Westhope, North Dakota. The 31

Souris River spans roughly 574 km (357 mi) in in North Dakota (Owen et al., 1981).

Once entirely glaciated and covered by the glacial Lake Souris, the basin is primarily made of forested hills in the northeast, vast glacial plains in the east, and more hills in the southwest.

Vermillion River Basin (Missouri River Sub-drainage)

The Vermillion River basin located east of the Missouri River in southeastern

South Dakota drains roughly 4,267 km2 (1,647 mi2) of land (Figure 2). The basin is positioned in-between the James River basin to the west, and the Big Sioux River basin to the east. Once entirely glaciated, the river valley was formed roughly 12,000 years ago by the melting ice (Hoagstrom et al., 2007; Christensen and Stephens, 1967). The East and

West Forks of the Vermillion River begin at the headwaters of the basin near Cherry

Lake, Lake Preston, Lake Thompson, Lake Whitewood, and Spirit Lake west of

Brookings, South Dakota. The Little Vermillion Rivers headwaters begin between

Howard and Madison, South Dakota, where it converges with the East Fork of the

Vermillion River near Montrose, South Dakota. The East and West Forks converge to form the Vermillion River just east of Parker, South Dakota, where it continues to flow south until it converges with the Missouri River along the South Dakota-Nebraska border just south of Vermillion, South Dakota.

White River Basin (Missouri River Sub-drainage)

The White River basin located west of the Missouri River in south-central South

Dakota drains roughly 21,357 km2 (8,246 mi2) (Figure 2) (Hoagstrom et al., 2007). The basin is surrounded by the Bad River drainage to the north, the Cheyenne River drainage to the west, the Niobrara River drainage in the south, and the Missouri River drainage to 32 the east. Much of the White River basin receives runoff from the western end of the

Badlands, which contributes to the high turbidity of the major river within the basin, the

White River. The White River headwaters are located just east of Harrison, Nebraska in the very northwest corner of the state. The White River flows northeast until it crosses the

Nebraska-South Dakota border just west of Pine Ridge, South Dakota, where it continues to flow northeast along the Badlands and eventually empties into Lake Oahe south of

Oacoma, South Dakota.

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Benson, R.D. 1983. A preliminary assessment of the hydrologic characteristics of the

James River in South Dakota. U.S. Geological Survey Water Resources

Investigations Report, Pierre, South Dakota, U.S.

Billesbach, D.P., and T.J. Arkebauer. 2012. First long-term, direct measurements of

evapotranspiration and surface water balance in the Nebraska Sand Hills.

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processes in a northern mixed-grass prairie, USA. Ecological Applications

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Bleed, A.S., and C.A. Flowerday. 1998. An atlas of the sand hills conservation and

survey division. University of Nebraska, Lincoln, Nebraska.

Bryce, S.A., J.M. Omernik, D.A. Pater, M. Ulmer, J. Schaar, J. Freeouf, R. Johnson, P.

Kuck, and S.H. Azevedo. 1996. Ecoregions of North and South Dakota, (color

poster with map, descriptive text, summary tables, and photographs): Reston

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Chapman, S.S., J.M. Omernik, J.A. Freeouf, D.G. Huggins, J.R. McCauley, C.C.

Freeman, G. Steinauer, R.T. Angelo, and R.L. Schlepp. 2001. Ecoregions of

Nebraska and Kansas (color poster with map, descriptive text, summary tables

and photographs): Reston, Virginia, U.S. Geological Survey (map scale

1:1,950,000).

Christensen, C., and J. Stephens. 1967. Geology and hydrology of Clay County, South

Dakota. South Dakota Geological Survey Bulletin 19, Pierre.

DeWitt, E., J.A. Redden, D. Buscher, and A.B. Wilson. 1989. Geologic map of the Black

Hills area, South Dakota and Wyoming: U.S. Geological Survey Miscellaneous

Investigations Series Map I-1910, I sheet, scale 1:250,000.

Duehr, J.P. 2004. Fish and habitat relations at multiple spatial scales in Cheyenne River

basin, South Dakota. M.S. Thesis, South Dakota State University, Brookings,

South Dakota.

Galat, D.L., C.R. Berry, W.M. Gardner, J.C. Henderson, G.E. Mestl, G.J. Power, C.

Stone, and M. R. Winston. 2005. Spatiotemporal patterns and changes in Missouri

River fishes. American Fisheries Society Symposium 45:249-291. 34

Goolsby, D.A., W.A. Battaglin, G.B. Lawrence, R.S. Artz, B.T. Aulenbach, R.P. Hooper,

D.R. Keeny, and G.J. Stensland. 1999. Flux and Sources of Nutrients in the

Mississippi-Atchafalaya River Basin: Topic 3 Report for the Integrated

Assessment on Hypoxia in the Gulf of Mexico. NOAA Coastal Ocean Program

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MD. 130pp.

Gonzalez, M.A. 2003. Continental Divides in North Dakota and North America. North

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Griffith, G. 2010. Level III North American Terrestrial Ecoregions: United States

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Cooperation.

Hendon, D., and J.A. Matthews. 2014. Pleistocene. Encyclopedia of Environmental

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Herman, G.S., and L.A. Johnson. 2007. Geology, Geography, and Climate. State

Historical Society of North Dakota. 112pp.

Hoagstrom, C.W., S.S. Wall, J.G. Kral, and B.G. Blackwell. 2007. Recent Zoogeography

of South Dakota fishes. Pages 37-89 in C. Berry, K. Higgins, D. Willis, and S.

Chipps, editors. History of fisheries and fishing in South Dakota. South Dakota

Department of Game, Fish and Parks, Pierre.

Horton, A.H., and R. Follansbee. 1906. Surface water supply of upper Mississippi river

and Hudson Bay drainages. Department of the Interior United States Geological

Survey, 1907. 35

Krause, J.R., K.N. Bertrand, A. Kafle, and N.H. Troelstrup, Jr. 2013. A fish index of

biotic integrity for South Dakota’s Northern Glaciated Plains Ecoregion.

Ecological Indications 34:313-322.

Maderak, M.L. 1966. Sedimentation and chemical quality of surface water in the Heart

River drainage basin, North Dakota. United States Government Printing Office,

Washington.

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richness in rivers. Ecography 18:345-352.

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South Dakota basins affected by the Garrison Diversion Unit. Fisheries Research

Unit, University of North Dakota, Grand Forks.

Pegg, M.A., C.L. Pierce, and A. Roy. 2003. Hydrological alteration along the Missouri

River Basin: a time series approach. Aquatic Sciences 65:63-72.

Pirner, S.M. 2018. The 2018 South Dakota integrated report for surface water quality

assessment. South Dakota Department of Environment and Natural Resources.

206pp.

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mining in the Black Hills, South Dakota, USA. Environmental Geology 27:38-53.

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sucker populations in the Black Hills, and an assessment of their thermal 36

tolerance. M.S. Thesis. Department of Wildlife and Fisheries Sciences, South

Dakota State University, Brooking, South Dakota. 116pp.

Shearer, J.S., and C.R. Berry, Jr. 2002. Index of biotic integrity utility for the fishery of

the James River of the Dakotas. Journal of Freshwater Ecology 17:575-588.

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threatens grasslands and wetlands. Proceedings of the National Academy of

Sciences 110:4134-4139.

38

CHAPTER 2

HISTORY OF ICHTHYOLOGY IN THE DAKOTAS

Prehistoric Fishes Evidence of both freshwater and marine fishes present in the Dakotas region well before humans inhabited earth comes from fossils scattered throughout North Dakota and

South Dakota. The Pleistocene Epoch (The Great Ice Age), or the last continental glaciation period, covered much of the eastern Dakotas, resulting in few fossil deposits in the eastern Dakotas. However, the western half of the Dakotas was primarily unglaciated, and fossils are very common in exposed rock. In fact, the oldest fish in the world

(Anatolepis) was found within the Black Hills, in the Bear Lodge Mountains of eastern

Wyoming, near the border of South Dakota (Parris et al., 2007). Fossils of the group of sharks known as bradyodonts, have also been discovered within the Black Hills region in rock formations over 300 million years old (Parris et al., 2007). These fossils are primarily teeth, as sharks within the class of vertebrates called Chondrichthyes, have bodies made of cartilage and do not preserve well in fossil form. Rock quarries in northeastern South Dakota, specifically Grant County, have also revealed fossils of sharks (Parris et al., 2007). Many of the fossils found throughout the western part of the

Dakotas are from bony fishes, of the vertebrate class Osteichthyes, and are found within

Paleozoic rock.

1800’s The very first descriptions of fishes from the Dakotas were written in the expedition journals of Meriwether Lewis and William Clark in 1804, where they recorded using fish as a food source. One of their journal entries notes a “white 39 catfish…with its eyes small and a tail much like that of a dolphin”. This description was proposed to be what we now call a Channel Catfish, Ictalurus punctatus. Between 1853 and 1855, a physician named Dr. John Evans made the first official scientific recordings of fish species within the Dakotas. Dr. Evans was sent out by the United States government to conduct surveys for the Pacific Railroad in the Dakotas region during the westward expansion. His findings out of Fort Pierre, South Dakota included Goldeye

(Hiodon alosoides), Shorthead Redhorse (Moxostoma macrolepidotum), Western Silvery

Minnow (Hybognathus argyritis), Flathead Chub (Platygobio gracillis), Creek Chub

(Semotilus atromaculatus), Channel Catfish (Ictalurus punctatus), River Carpsucker

(Carpiodes carpio), and Paddlefish (Polyodon spathula). These collections were then sent by Dr. John Evans to Charles Girard, who officially published the findings (Girard,

1856; Girard, 1858) and established the first species list for the Dakotas, making them the oldest known specimens of the Dakotas.

During the same time frame, Robert Kennicott while working for the Smithsonian

Institution, was recording some new discoveries of Johnny Darter (Etheostoma nigrum) from the Goose River and Blackside Darter (Percina maculata) from the Maple River in

North Dakota. This then increased the number of known species from the Dakotas region to ten by 1857. In 1859, Girard published the first known collections of Iowa Darters

(Etheostoma exile) from North Dakota, where he listed a collection made in the Little

Muddy River by Dr. George Suckley, and another collection in the Cannonball River by

Dr. F.V. Hayden (Girard, 1859).

In 1873 and 1874, Dr. Elliot Coues made the next reports of fishes collected in the

Dakotas as he was traveling west to Montana for the United States Geological and 40

Geographical Survey of the Territories. Dr. Coues sent his new discoveries off to one of the most well-known ichthyologists to date, David Starr Jordan, who published the

Report on the Collection of Fishes Made by Dr. Elliot Coues, U.S.A. in Dakota and

Montana during the Seasons of 1873 and 1874 (Jordan, 1878) and added the first records of Shovelnose Sturgeon (Scaphirhynchus platorhynchus) and Northern Pike (Esox lucius) from the Dakotas. Jordan also described the Lake Chub (Couesius plumbeus), from Dr.

Coues collections, and named it after him.

Edward D. Cope made the next documentation of existing species in 1876. While on a journey to collect fossils from the Moreau River, he sampled and collected fishes from Battle Creek, a tributary of the Missouri River. This sampling site has also been documented as Blue Blanket Creek (Bailey and Allum, 1962) and LeBeau Creek (Berry et al., 2007). During his journey, Cope was the first to document the presence of Burbot

(Lota lota), Longnose Dace (Rhinichthys cataractae), Northern Redbelly Dace

(Chrosomus eos), Longnose Gar (Lepisosteus osseus), and Shortnose Gar (Lepisosteus platostomus) in the Dakotas (Cope, 1879).

The Dakotas region officially became the separate states, North Dakota and

South Dakota in 1889. The first survey documented after this separation was made by

Seth E. Meek in southeastern South Dakota in 1891 where he collected fishes between

Sioux City, Iowa and Sioux Falls, South Dakota in the Missouri and Big Sioux rivers

(Meek, 1892). This survey served to be important for the state of South Dakota because it revealed 24 new species of fish not yet previously reported in the Dakotas.

In 1892, Ulysses S. Cox and Albert Jefferson Woolman conducted stream surveys in western Minnesota and eastern North Dakota where they found a total of six, non- 41 previously reported species from both North Dakota and South Dakota. Specifically, the area surveyed included streams in the Minnesota, James, and Red River of the North watersheds, as well as Lake Traverse and Big Stone Lake (Woolman, 1896). Of these collections was the first known Pugnose Shiner (Notropis anogenus) from North Dakota in the Sheyenne River, but was misidentified by Woolman as Bleeding Shiner, Notropis zonatus. Carl Hubbs at the University of Michigan Museum of Zoology later corrected

Woolmans misidentification (UMMZ, catalog number: 61945).

In 1893 while assessing the potential of a trout fishery, the ichthyofauna of the

Black Hills streams were surveyed by Barton Warren Evermann (Evermann, 1892). In

1896, Evermann and Ulysses Orange Cox co-wrote and published the Report Upon the

Fishes of the Missouri River Basin, which compiled official species lists for each of the states within the Missouri River drainage including North and South Dakota (Everman and Cox, 1896). This report was also one of the first publications to summarize all of the previous ichthyofauna surveys in both North Dakota and South Dakota and added another

13 species of fish to the Dakotas.

Early 1900’s

In 1929, the first Fishes of North Dakota was published by Thomas Leroy

Hankinson and included a total of 57 species for the state, 13 of which were new to the species list of North Dakota (Hankinson, 1929). This publication summarized all historical collections made in North Dakota to date and included collections that had been made statewide between 1921 and 1928 by the University of North Dakota Biological

Station (UNDBS) and the Museum of Zoology at the University of Michigan (UMMZ).

Collectors from the UNDBS included R.T. Young, Robert Johnstone, and Hankinson 42 himself. Collectors from UMMZ included Carl Hubbs, Leonard P. Schultz, and Crystal

Thompson.

In 1927, the South Dakota Department of Game and Fish (now South Dakota

Game, Fish and Parks) funded William Over and Edward Churchill from the University of South Dakota, to conduct lake surveys throughout eastern South Dakota (Churchill and

Over, 1927). In 1927 and 1928, the project was expanded to a statewide survey of all streams and lakes in South Dakota. From their findings, Churchill and Over published the first Fishes of South Dakota out of the University of South Dakota, which was the first fishes book dedicated to the state of South Dakota and included a dichotomous key to the family level (Churchill and Over, 1933). This publication included a total of 81 species,

12 of which were new to the species list for South Dakota, and provided brief summaries of each species, and even a few species illustrations.

The 1930’s proved to be a rough time for our nation’s history. During this era, a period of severe dust storms known as the Dust Bowl, swept through the American and

Canadian prairies and severely interrupting the landscapes ecology and agriculture. Many of the lakes and ponds within North and South Dakota dried up during this time period, and as a result, numerous stocking events took place once water levels naturally recovered overtime. Also, during this time period, the United States was undergoing the

Great Depression. President Franklin D. Roosevelt granted the National Industrial

Recovery Act of 1933, which authorized for the construction of Fort Peck Dam along the

Missouri River in northeastern Montana. Completed in 1940 and constructed by the

United States Army Corps of Engineers as the first dam along the Missouri River, Fort

Peck Dam impounds Fort Peck Lake, the nation’s fifth largest reservoir totaling 216 km 43

(134 mi) in length. Multiple problems with flooding in the lower portions of the Missouri

River drainage, a lack of water for irrigation purposes, and the success of Fort Peck Dam encouraged the United States government to pass the Flood Control Act of 1944, which resulted in the Pick-Sloan Plan. The Pick-Sloan Plan was plan for the United States Army

Corps of Engineers to construct five more major dams along the Missouri River between

Riverdale, North Dakota and Sioux City, Iowa. These dams and their adjacent reservoirs would help reduce flood damage, be available for irrigational use and water supply, generate electricity, be available for recreational use, fish and wildlife benefits, navigation, and create jobs (CMRES, 2002).

Post Reservior Construction: 1950’s to Today

In 1962, a revised edition of Fishes of South Dakota, was published by Reeve M.

Bailey and Marvin O. Allum and added recent surveys and historical data from across the state (Bailey and Allum, 1962). The revised publication now included a total of 110 species of fish from the state and included survey information from pre- and post- reservoir construction on the Upper Missouri River. Also included were more detailed summaries of each species, a dichotomous key, and ten new species to the species list for the Dakotas.

Although North Dakota did have some publications which provided a species list, until 1981 no real “Fishes of” book had been published for the state until J.B. Owen published the Distribution of Fishes in North and South Dakota basins affected by the

Garrison Diversion Unit (Owen et al., 1981). Although the book only focuses on the

James, Souris, Sheyenne and Wild Rice Rivers, it was one of the more complete and in- depth publications on fishes from North and South Dakota, as it included a dichotomous 44 key (written by Dr. Steve Kelesch, Professor and University of North Dakota), species summaries, distribution maps, and added 10 new species to the fish list for the Dakotas.

Since Bailey and Allum published the Fishes of South Dakota in 1962, no new books have been published to include the updated knowledge gained throughout the last five decades of fish surveys in either North or South Dakota. In 1994, the Fishes of North

Dakota booklet was published by the North Dakota Game and Fish Department, and included 36 species. However, this still is not considered a true “Fishes of” book, as it was written as a short outreach publication for the general public, due to its lack of distribution maps and a dichotomous key. The more detailed South Dakota equivalent was published two different times by South Dakota Game Fish and Parks and South

Dakota State University. In 1994, Robert M. Neuman and David W. Willis published The

Guide to the Common Fishes of South Dakota (Neuman and Willis, 1994) and in 2011, the equivalent of a second edition was published by Christopher W. Hoagstrom. The later edition lists 125 species for the state, however illustrations, life history summaries, and distribution maps are only included for 53 species. Although distribution maps are included, these maps show a rather broad range at the river system level and are not point specific.

Aside from state ichthyological publications, numerous individuals from state and federal agencies as well as universities have dedicated their time to contributing to the ichthyology of the Dakotas by sampling, recording, and collecting all types of fisheries data for various studies and projects. This data was carefully compiled and used for this project to create a fisheries distribution database. This distribution database is a conglomeration of over 400,000 records of fishes sampled from the 1800’s to today in the 45

Dakotas. From this database came the species distribution maps for each of the fish species found within North Dakota and South Dakota. These maps are not included within this thesis; however, they are forthcoming in the publication of the Fishes of the

Dakotas book. By synthesizing historical ichthyological data from both North Dakota and

South Dakota, we have been able to add and update species literature and documentation to serve as a valuable tool for students, researchers, biologists, and enthusiasts.

Literature Cited:

Churchill, E.P., and W.H. Over. 1933. Fishes of South Dakota. South Dakota Department

of Game and Fish, Pierre, 87.

Everman, B.W., and U.O. Cox. 1896. Report upon the fishes of the Missouri River basin.

U.S. Government Printing Office, 1896.

Jordan, D.S. 1878. A catalogue of the fishes of the fresh waters of North America.

Author’s ed. Washington: Department of the Interior.

Parris, D.C., G.A. Bishop, and K.F. Higgins. 2007. Pages 19-36 in C. Berry, K. Higgins,

D. Willis, and S. Chipps, editors. History of fisheries and fishing in South Dakota.

South Dakota Department of Game, Fish and Parks, Pierre.

46

CHAPTER 3

FAMILY PETROMYZONTIDAE

Introduction

Lampreys are one of the most ancient and unique groups of fishes found in the

Dakotas. The Lamprey family, Petromyzontidae, consists of 42 species in eight genera.

Two species, the Chestnut Lamprey, Icthtyomyzon castaneus, and the Silver Lamprey,

Icthtyomyzon unicuspis, are native species to the Dakotas. All species of lampreys undergo metamorphosis between two main life stages: larvae, called ammocoetes, and adulthood. Ammocoetes have a toothless hood-shaped mouth, and eyes that are covered by skin, making them completely blind. Ammocoetes burrow into sandbars, soft substrates, or beds of organic debris where they filter feed by sticking their heads out of their burrows to catch drifting plankton and bacteria. After a number of years, with the amount of time varying by species, ammocoetes will undergo metamorphosis, and develop functioning eyes, seven-gill openings on each lateral side, and a round or oval shaped sectorial mouth with rows of teeth and fringed papillae. Adult lamprey also have entirely cartilaginous, long, slender, snake-like bodies with no scales and have a single dorsal fin continuous with the caudal fin, and no anal, pelvic, pectoral fins. Lampreys in the Dakotas may often times be confused with the American Eel, Anguilla rostrata, from the family Anguillidae, however, eels can be easily distinguished by the presence of jaws and pectoral fins.

Members of Petromyzontidae are only found across the Northern Hemisphere

(North America, Asia and Europe) in cooler waters, simply because ammocoetes are 47 unable to survive in warmer water temperatures. Lamprey species can either be anadromous, meaning that they move to marine waters after they undergo metamorphosis in freshwater, or they can be entirely freshwater species, like the Chestnut and Silver

Lamprey. Both the Chestnut and Silver Lamprey are native to the Hudson Bay, St.

Lawrence-Great Lakes and Mississippi River drainages.

Each species of lamprey can be separated into two groups: parasitic and non- parasitic species. Parasitic adult lamprey attach to hosts (mainly other ray-finned fishes) after metamorphosis, and feed on their blood, tissue and other fluids using their sectorial disc. This parasitic behavior often times does not kill the host, however it likely weakens the hosts immunity to disease and bacteria. On the contrary, non-parasitic adult lamprey do not feed as adults after they complete metamorphosis. The Chestnut and Silver

Lamprey are both parasitic. After feeding for one to two years, both the Chestnut and

Silver Lamprey migrate upstream to the spawning grounds to reproduce. All species of lamprey spawn during the spring and summer, with the Chestnut and Silver Lamprey beginning their upstream migrations to the spawning areas in the late winter and early spring. Both Chestnut and Silver Lamprey prefer to spawn in shallow reaches of smaller, low order streams with moderate current over sand or gravel substrate. Spawning behavior of lamprey is very similar for all species. Female lampreys use their mouth to stabilize themselves onto a hard and sturdy surface within or over a nest, while the males attach themselves to the head of the female and wrap their bodies around the females body. During this time, their bodies vibrate together to release eggs and sperm. Like all species of lamprey, Chestnut and Silver Lamprey are semelparous, meaning that they die, or end their life cycle after a single spawning event. 48

Chestnut Lamprey, Ichthyomyzon castaneus (Girard, 1858)

Etymology and Synonyms: Ichthyomyzon = “sucker of fish”; castaneus = “a chestnut”, referring to the adult coloration.

Description: Chestnut Lamprey have an elongate, cylindrical, laterally compressed body posteriorly. Dorsally gray, tan, to olive; laterally tan to olive, sometimes mottled; ventrally light brown to light yellow-cream; dorsal fin with brown tint. Head small, cylindrical. Eye small, placed dorsolaterally on head. Jaw absent. Mouth disk or oval shaped outlined with fringed papillae; maximum diameter greater than maximum diameter of body in adults. Teeth present; circumorals bicuspid (double pointed), sharp, claw-like; 19-25 teeth in circumoral row; 3-6 teeth in anterior row; 7-11 teeth in lateral rows; supraoral cusps 2-5, close together. Gill openings 7, small and pore-like on each side. Dorsal fin elongate with a shallow notch lacking spines or rays, connected to a protocercal caudal fin, and continuous into the anal fin. No paired fins present. Scaleless.

Lateral sides with 50-56 myomeres, between the last gill opening and vent. Lateral pores small, pigmented in ammocoetes and adults. Young larvae darker tan to brown in color without eyes or a sucking disk, instead displaying a hood-shaped mouth without fringed papillae.

Similar Species: Closely resembles the Silver Lamprey. Silver Lamprey more gray in color, have unicuspid (single pointed) circumoral teeth, 46-53 myomeres on lateral sides, and adults with pigmented lateral pores. American Eel have a snake-like body with jaws and pectoral fins present.

Distribution and Habitat: Native to southern Canada and the upper mid-western United

States from the Red River basin in eastern North Dakota and western Minnesota in the 49 west, east through the St. Lawrence-Great Lakes, upper Mississippi, and Ohio River drainages, and south to the Gulf of Mexico in Louisiana and Texas.9 Occurs only within the Red River basin of North Dakota. Reported to occur in the Bios de Sioux River in

South Dakota. Occurs more frequently in smaller waterbodies than the Silver Lamprey.1

Adults found within medium-sized streams, large rivers, and impoundments with moderate flow over gravel, sand, or silt substrate.1,6,10 Ammocoetes require areas with sand and silt substrate, beds of organic matter, and light vegetation such as Chara, in backwaters with moderate current.11

Reproduction: Spawning migrations begin in late winter and early spring and consist of upstream movements to shallow reaches with moderate current in low order streams with sand or gravel substrate. Spawning date varies with latitude, with preferred spawning water temperature less variable.5 Spawning aggregations in Wisconsin take place May to

June, and have been observed in water 15-22°C (59-71.6°F) at depths of 18-46 cm (0.59-

1.51 ft.).5 Females use their mouth to suction onto a hard surface, such as a rock, and males wrap themselves around the females body to instigate a quivering motion to release eggs.2 Adults construct nests by moving stones with their disk-shaped mouths, and excavating a depression roughly 60 cm (23.6 in) wide, 100 cm (39.4 in) long, and 5 cm

(1.97 in) deep, within gravel or sand substrate.2 As many as 50 Chestnut Lampreys have been observed spawning in the same nest.2 No parental care is given to eggs or young.

Fecundity increases with the length of female. Estimated fecundity from Michigan

10,144-18,563 eggs per female.12 Average fecundity from a 255 mm (10.04 in) TL adult female in its second year in Wisconsin reported as roughly 13,000 eggs 0.6-0.7 mm

(0.02-0.03 in) in diameter.1 Death occurs shortly after spawning. After hatching, larvae 50 will drift downstream eventually burrowing into sand bars or beds of organic debris where they will remain for 5-7 years.11 During this period they are referred to as ammocoetes. Metamorphosis into the adult form begins in late summer, and by springtime they emerge from their burrows to seek out a host.7

Age and Growth: Little information is available on age and growth. Newly morphed adults emerging from their burrows roughly 100 mm (3.94 in) TL.7 During the parasitic phase, most growth takes place during summer months.5 Females tend to achieve greater lengths than males.5 Longevity presumed to be 6-7 years. Capable of reaching 355.6 mm

(14 in) TL.5,8,12

Food and Feeding: Ammocoetes filter feed by sticking their heads out of their burrows to catch drifting prey such as diatoms, phytoplankton, detritus, and bacteria.1 Adults are parasitic and attach themselves in the springtime to hosts, by cutting through scales with their teeth, and extracting blood mainly on the dorsal side of the host.3,5,7 Majority of feeding takes place May to October, with a peak in July followed by a period of feeding inactivity in winter.5,7 Parasitic adults tend to be size selective and are strongly associated with relatively large host species with small scales or naked skin such as Paddlefish, Pike,

Catfish, and Redhorse species.4,5 Adults are parasitic for 1-2 years before migrating to spawning grounds and ending their lifecycle.7

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Case, B. 1970. Spawning behavior of the Chestnut Lamprey (Ichthyomyzon

castaneus). Journal of the Fisheries Research Board of Canada 27:1872-1874. 51

3. Cochran PA. 1986a. The daily timing of lamprey attacks. Environmental Biology

of Fishes. 16:325-329.

4. Cochran, P.A. 1994. Why lampreys and humans “compete” (and when they

don’t): toward a theory of host species selection by parasitic lampreys. Pages 329-

345 in Stouder, D.J., K.L. Fresh, and R.J. Feller (editors). Theory and application

in fish feeding ecology. Belle W. Baruch Library in Marine Science, No. 18,

University of South Carolina Press, Columbia, South Carolina.

5. Cochran, P.A. 2014. Field and laboratory observations on the ecology and

behavior of the Chestnut Lamprey Ichthyomyzon castaneus. Journal of Freshwater

Ecology 29:491-505.

6. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

7. Hall, J.D. 1963. An ecological study of the Chestnut Lamprey, Ichthyomyzon

castaneus (Girard), in the Manistee River, Michigan. PhD dissertation. University

of Michigan, Ann Arbor.

8. Moore, G.A., and M. Kernodle. 1965. A new size record for the Chestnut

Lamprey, Ichthyomyzon castaneus (Girard) in Oklahoma. Proceedings of the

Oklahoma Academy of Science 45:68-69.

9. Page, L.M., and B.M. Burr. 2011. Peterson field guide to freshwater fishes of

North America north of Mexico. 2nd ed. New York (NY): Houghton Mifflin.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 52

11. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

12. Starrett, W.C., W.J. Harth, and P.W. Smith. 1960. Parasitic lampreys of the genus

Ichthyomyzon in the rivers of Illinois. Copeia 1960:337-346.

53

Silver Lamprey, Ichthyomyzon unicuspis (Hubbs & Trautman, 1937)

Etymology and Synonyms: Ichthyomyzon = “sucker of fish”; unicuspis = “unicuspid”, referring to the circumoral teeth.

Description: Silver Lamprey have an elongate, cylindrical, laterally compressed body posteriorly. Dorsally dusky blue to dark gray; laterally bluish gray; ventrally dark gray; dorsal fin may have a brown tint, fading to gray on lateral and ventral sides. Head small, cylindrical. Eye small, placed dorsolaterally on head. Jaw absent. Mouth disk or oval shaped outlined with fringed papillae; maximum diameter greater than maximum diameter of body in adults. Teeth present; circumorals unicuspid (single point), sharp, claw-like; 18-25 teeth in circumoral row (in rare cases, they may possess as many as 2 bicuspid circumorals); 2-4 teeth in anterior row; 4-9 teeth in lateral rows; supraoral cusps

1-4, close together. Gill openings 7, small and pore-like on each side. Dorsal fin elongate with a shallow notch lacking spines or rays, connected to a protocercal caudal fin, and continuous into the anal fin. No paired fins present. Scaleless. Lateral sides with 46-53 myomeres, between the last gill opening and vent. Lateral pores small, unpigmented in ammocoetes; pigmented in large adults. Young larvae without eyes or sucking disk instead displaying a hood-shaped mouth without fringed papillae.

Similar Species: Closely resembles the Chestnut Lamprey. Chestnut Lamprey more tan to brown in color, have bicuspid (double pointed) circumoral teeth, 50-56 myomeres on lateral sides, and adults with unpigmented lateral pores. American Eel have a snake-like body with jaws and pectoral fins present.

Distribution and Habitat: Native to the Red River basin in eastern North Dakota and western Minnesota in the west, east through the St. Lawrence-Great Lakes, upper 54

Mississippi, and Ohio River drainages to New York, and south to central Tennessee.

Occurs outside its native range due to canal connection. Occurs within the Red River basin in North Dakota, and the Missouri River basin in South Dakota near Lewis and

Clark Lake. Adults found within mainstems of large rivers, streams, and tributaries as well as small to large lakes or impoundments with gravel substrate. Ammocoetes require sand bars or beds of organic debris within muddy pools or backwaters.

Reproduction: Information on reproductive behavior is scarce. Spawning migrations begin when water temperature reaches roughly 10°C (50°F), and consist of movements made upstream to areas near run to riffle-like habitat with gravel, cobble, and boulder substrate.1,3 Males arrive at the spawning grounds prior to females, and may attract females with the release of sexual pheromones.4,5 Spawning aggregations take place

May-June, and have been observed in clear to turbid waters 12.8-22.8°C (55.04-73.04°F) and depths of 23-79 cm (0.75-2.59 ft.).3,4,8 As many as 10 Silver Lampreys have been observed spawning in the same nest.8 Nests roughly 30-122 cm (11.81-48.03 in) in diameter and 2.5-15 cm (0.98-5.91 in) deep, and are constructed in sand or gravel substrate in streams of moderate gradient near riffles at depths roughly 8-15 cm (3.15-

5.91 ft).8,10,13 No parental care is given. Fecundity increases with length of female.4,9

Average fecundity from a 247 mm (9.72 in) TL female reported as 19,012 eggs, and a

326 mm (12.83 in) TL female produced roughly 27,400 eggs 0.8-1.0 mm (0.03-0.04 in) in diameter.1,7 Death occurs shortly after spawning. Larvae hatch within 5 days in water

18.4°C (65.12°F), and will drift downstream, eventually burrowing into sand bars or beds of organic debris where they will remain for 4-7 years.1,10,11 During this period, they are 55 referred to as ammocoetes. Metamorphosis into the adult form begins in August, and by springtime they emerge from their burrows to seek out a host.1

Age and Growth: Little information is available on age and growth. Newly morphed adults emerging from their burrows roughly 89-152 mm (3.50-5.98 in) TL.1 Growth mainly occurs during the summer months in the parasitic phase.4 Females tend to achieve greater lengths than males.4 Longevity 6-7 years.4 Capable of reaching 395 mm (15.55 in)

TL.1,12

Food and Feeding: Ammocoetes filter feed by sticking their heads out of their burrows to catch drifting prey such as algae, pollen, diatoms, and protozoans.1,6 Adults are parasitic and attach themselves to hosts by cutting through scales with their teeth, and extracting blood. Adults also rely on food they have acquired as ammocoetes. Lampreys tend to be size selective with their choice of host.2,3 Parasitic adults strongly associated with relatively large host species with small scales or naked skin such as Paddlefish, Lake

Trout, Catfish, and Sturgeon species.1,8 Although a large and numerous species, Common

Carp are often not selected as hosts due to the Silver Lampreys inability to puncture their large, heavy scales.3 Adults attach to their hosts in the springtime, and are mainly seen attached on the dorsal side of the host.3 Adults are parasitic for roughly one year before migrating to spawning grounds and ending their lifecycle.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Cochran, P.A. 1994. Why lampreys and humans “compete” (and when they

don’t): toward a theory of host species selection by parasitic lampreys. Pages 329-

345 in Stouder, D.J., K.L. Fresh, and R.J. Feller (editors). Theory and application 56

in fish feeding ecology. Belle W. Baruch Library in Marine Science, No. 18,

University of South Carolina Press, Columbia, South Carolina.

3. Cochran, P.A., and J. Lyons. 2004. Field and laboratory observation on the

ecology and behavior of the Silver Lamprey (Ichthyomyzon unicuspis) in

Wisconsin. Journal of Freshwater Ecology 19:245-253.

4. Cochran, P.A., and J.E. Marks. 1995. Biology of the Silver Lamprey,

Ichthyomyzon unicuspis, in Green Bay and the lower Fox River, with a

comparison to the Sea Lamprey, Petromyzon marinus. Copeia 1995:409-421.

5. Hardisty, M.W. 1986. General introduction to lampreys. Pages 19-83 in Holčík, J.

(editor). The freshwater fishes of Europe. Vol. 1, Part I, Petromyzontiformes.

AULA-Verlag, Weisbaden, Germany.

6. Harlan, J.R., and E.B. Speaker. 1951. Iowa Fish and Fishing. State Cons. Comm.

237p.

7. Manion, P.J., and L.H. Hanson. 1980. Spawning behavior and fecundity of

lampreys from the upper three Great Lakes. Canadian Journal of Fisheries and

Aquatic Sciences 37:1635-1640.

8. Morman, R.H. 1979. Distribution and ecology of lampreys in the Lower

Peninsula of Michigan, 1957-75. Great Lakes Fishery Commission Technical

Report No. 33.

9. Schuldt, R.J., M.F. Fodale, and W.J. Johnson. 1987. Prespawning characteristics

of lampreys native to Lake Michigan. Journal of Great Lakes Research 13:264-

271. 57

10. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

11. Smith, A.J., J.H. Howell, and GW. Piavis. 1968. Comparative embryology of five

species of lampreys of the upper Great Lakes. Copeia 1968:461-469.

12. Starrett, W.C., W.J. Harth, and P.W. Smith. 1960. Parasitic lampreys of the genus

Ichthyomyzon in the rivers of Illinois. Copeia 1960:337-346.

13. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press. 683p.

58

CHAPTER 4

FAMILY ACIPENSERIDAE

Introduction

The Sturgeon family, Acipenseridae, consists of four genera that comprise a total of 25 extant species. The majority of the 25 species occur throughout Europe and Asia, and a total of eight occur within North America (five species within the genera

Acipenser, and three within the genera Scaphirhynchus) along both the Atlantic and

Pacific coasts, as well as inland freshwaters. Sturgeon are a primitive group of fishes that evolved during the end of the Triassic period nearly 200-250 million years ago, making them the most ancient extant group of the bony fishes (Actinopterygii). Sturgeons are the largest fishes found in freshwater, and display a fusiform, elongate, robust, and relatively dorsoventrally flattened body shape with a short dorsal fin set far posterior on the body and a distinguishable heterocercal tail. They also possess two sets of barbels located inferior on the head, anterior to the subterminal mouth. Sturgeons bodies are covered in five rows of bony plates, called scutes, that are situated dorsally, laterally, and ventrally.

In North America, sturgeon inhabit the benthic habitats of fresh, brackish, and marine waters. While cruising just above the bottom substrate, they use their sensory barbels with taste buds to detect food. Carvinorous during all stages of life, sturgeon begin to feed on zooplankton during their larval stage before undergoing an ontogenetic diet shift as they mature as juveniles and adults, and eventually consume benthic invertebrates and smaller fishes. Although some species of sturgeon spend the majority of their lifetimes in brackish or marine waters, all sturgeon migrate and spawn in freshwater 59 rivers. The largest species of North American sturgeon, the White Sturgeon, Acipenser transmontanus, is capable of migrating more than 1,000km (621mi) from the ocean to freshwater to spawn, whereas the smallest of the North American Sturgeon, the Shortnose

Sturgeon, Acipenser brevirostrum, typically migrates less than 200km (124mi).

Although often thought of as one of the more admirable, historical groups of fishes, sturgeons have been experiencing dramatic declines in populations due to several biological factors coupled with anthropogenic disturbances. Sturgeon are a naturally slow growing group of fishes that experience delayed maturation, and often do not become sexually mature for several years, sometimes even decades. Populations of more northern species of sturgeon often times mature later than southern species of sturgeon, and males frequently reach sexual maturity before females. Spawning events are also infrequent, and often have several years in-between them. For example, female Pallid Sturgeon,

Scaphirhynchus albus, spawn once every 4-10 years, while males spawn once every 2-3 years. Slow growth, delayed maturation, and infrequent spawning already present somewhat of a challenge for sturgeon populations to persist over time, which they have proven to do so. However, habitat alterations such as the construction of dams are one of the several anthropogenic disturbances that negatively affect the life history of sturgeon.

Three species of sturgeon occur within the Dakotas: the Lake Sturgeon, Acipenser fulvescens, Shovelnose Sturgeon, Scaphirhynchus platorhynchus, and the Pallid

Sturegon. Pallid Sturgeon are currently listed as federally endangered.

60

Lake Sturgeon, Acipenser fulvescens (Rafinesque, 1817)

Etymology and Synonyms: Acipenser = Latin for sturgeon, derived from akis, meaning

“point”, and pente, meaning “five”, referring to five rows of body scutes; fulvescens =

“yellowish brown”, referring to the coloration on the dorsal and lateral sides.

Description: Body fusiform but dorsoventrally flattened. Dorsally dark olive to gray; laterally yellow-brown to gray; ventrally white; body with five rows of large scutes; dorsal scutes 9-17; lateral scutes 29-42; ventral scutes 7-12. Head elongate, dorsoventrally flattened. Snout short, rounded, conical, slightly upturned. Eye small, placed dorsolaterally on head; spiracle present, slightly posterior of eye. Mouth inferior.

Lips fleshy, lower lip with two papillose lobes. Barbels present, smooth; four positioned anterior of mouth on ventral side of the snout. Frenum absent. Gill rakers 25-44. Dorsal fin set posterior on body with 35-40 rays. Adipose fin absent. Caudal peduncle short, thick; exposed portions of skin present between scutes. Caudal fin heterocercal with upper lobe more elongate than lower lobe, but not extending into a long filament. Anal fin with 25-30 rays. Pelvic fins positioned posterior on body, but insertions anterior to insertion of dorsal fin. Pectoral fins large, rounded. Lateral line complete, extending through upper lobe of caudal fin. Spawning adults similar to non-spawners. Juveniles similar to adults but with small, dark mottling on body.

Similar Species: Closely resembles the Pallid and Shovelnose Sturgeon. Pallid Sturgeon and Shovelnose Sturgeon have a more flattened and shovel shaped head, elongate and slender caudal peduncles without exposed areas of skin, lack spiracles, and have fringed barbels. 61

Distribution and Habitat: Native to eastern North America throughout the Great Lakes,

Hudson-James Bay, and Mississippi watersheds, and as far south as Alabama and northern Mississippi.1,12 Once abundant and widespread throughout its native range, populations have been declining due to anthropogenic disturbances such as pollution, commercial fishing, habitat fragmentation, and loss of spawning habitat.16 Occurs within the Red River of the North in North Dakota. Present in the mainstem of the Missouri

River below Gavin’s Point Dam in South Dakota, and historically occurred within Big

Stone Lake in the Minnesota River drainage. In 2014, a collaborative effort between

South Dakota Game, Fish and Parks and the Minnesota Department of Natural Resources was made to implement a stocking program in Big Stone Lake in hopes to restore what was once a native population. Inhabits large rivers and lakes with sand, gravel or mud substrate. Following hatching, larvae remain burrowed within the gravel substrate until the yolk sac is absorbed, after which they begin to drift downstream to nursery habitats which include shallow, low velocity areas with sand substrate and a predominance of dipteran larvae.4,16 Juveniles are often found within lower gradient streams at depths 0.5-

20 m (1.64-26.25 ft), over fine substrate.8,10,16 Adults frequently occupy deeper depths during fall and winter rather than during the spring and summer.6

Reproduction: Spawning migrations begin in early spring near ice-out, and consist of movements made upstream that are often over 129 km (80.16 mi), but are capable of exceeding 402 km (249.79 mi).1,18 Movement rates of Lake Sturgeon from the upper

Mississippi River system in the spring averaged roughly 0.50 km/day (0.31 mi/day).1,11

Spawning typically occurs between late April and early June, and takes place over shallow and swift shoals within moderate to high gradient rivers and streams over cobble, 62 gravel or rocky substrate.1,3,5,8 Males begin to move into spawning sites prior to females when temperatures reach 8.8-16.0°C (47.84-60.8°F).7,16 Females are known to wait until the water temperature reaches 19.1°C (66.38°F).7,16 Sexual maturation is delayed; mean age at sexual maturity of males from the Lake of the Woods and the Rainy River system reported as 16.8 years, and females 25.8 years.14 Males generally only spawn every 1-3 years, and females every 4-6 years, with the amount of time between spawning events increasing with latitude.1,13 Spawning behavior consists of several males swimming upstream alongside a single female, and vibrating their bodies against hers to initiate the release of eggs and milt.3 Fecundity highly variable, with females of the same size capable of producing 50,000-700,000 eggs.3 No nest is constructed and no parental care is given.5 Eggs black, demersal, adhesive to rocks, gravel or woody structures, and roughly 2.0-3.0 mm (0.08-0.12 in) in diameter.1,5 Hatching and incubation time dependent on water temperature, but occurs within 5-8 days in water 16-17°C (60.80-

62.60°F).5

Age and Growth: Larvae roughly 8 mm (0.31 in) long at hatching, and roughly 125 mm

(4.92 in) by their first fall.3,5 Mean lengths-at-age of individuals from Rainy Lake are reported as: age-15, 1165 mm (45.86 in) TL; age-20, 1233 mm (48.54 in) TL; age-25,

1273 mm (50.02 in) TL; age-30, 1334 mm (52.52 in) TL; age-35, 1442 mm (56.77 in)

TL; age-40, 1433 mm (56.42 in) TL.1 Females generally larger than males.1 Capable of reaching 2745 mm (9 ft).3 Longevity 154 years.9

Food and Feeding: Benthic, generalist, and opportunistic feeders.2,15,16 Forage by using their barbels to detect prey, and then capture prey by using their inferior mouth to suction feed. Lake Sturgeon diets are very diverse throughout its range, as they appear to be 63 advantageous and have plasticity.16 Diet items consist of a variety of benthic organisms including blood worms, mayfly nymphs, midge larvae, amphipods, gastropods, crayfish and other fishes such as Gizzard Shad.5,14,16,17,19

Literature Cited:

1. Adams, W.E. Jr. 2004. Lake sturgeon biology in Rainy Lake, Minnesota and

Ontario. M.S. Thesis, South Dakota State University, Brookings, South Dakota.

2. Beamish, F.W. H., D.L.G. Noakes, and A. Rossiter. 1998. Feeding ecology of

juvenile lake sturgeon, Acipenser fulvescens, in northern Ontario. Canadian Field

Naturalist 112:459-468.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Benson, A.C., T.M. Sutton, R.F. Elliott, and T.G. Meronek. 2005. Seasonal

movement patterns and habitat preferences of age-0 lake sturgeon in the lower

Peshtigo River, Wisconsin. Transactions of the American Fisheries Society

134:1400-1409.

5. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

6. Briggs, A.S., D.W. Hondorp, H.R. Quinlan, J.C. Boase, and L.C. Mohr. 2016.

Electric archival tags provide first glimpse of bathythermal habitat use by free-

ranging adult lake sturgeon Acipenser fulvescens. Journal of Freshwater Ecology

31:477-483.

7. Bruch, R.M., and F.P. Binkowski. 2002. Spawning behaviour of lake sturgeon

(Acipenser fulvescens). Journal of Applied Ichthyology 18:570-579. 64

8. Daugherty, D.J., T.M. Sutton, and R.F. Elliott. 2009. Suitability modeling of lake

sturgeon habitat in five northern Lake Michigan tributaries: implications for

population rehabilitation. Restoration Ecology 17:245-257.

9. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

10. Haxton, T.J. 2011. Depth selectivity and spatial distribution of juvenile lake

sturgeon in a large fragmented river. Journal of Applied Ichthyology 27:45-52.

11. Knights, B.C., J.M. Vallazza, S.J. Zigler, and M.R. Dewey. 2002. Habitat and

movement of lake sturgeon in the upper Mississippi River System, USA.

Transactions of the American Fisheries Society 131:507-522.

12. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

13. Magnin, E. 1966. Recherches sur les cycles de reproduction des sturgeons

Acipenser fulvescens Rafinesque, Annales de la Station Centrale d’Hydrobiologie

appliquee 9:8-242.

14. Mosindy, T., and J. Rusak. 1991. An assessment of lake sturgeon populations in

Lake of the Woods and the Rainy River 1987-90. Ontario Ministry of Natural

Resources, Lake of the Woods Fisheries Assessment Unit Report 1991:01,

Toronto.

15. Nilo, P. S. Tremblay, A. Bolon, J. Dodson, P. Dumont, and R. Fortin. 2006.

Feeding ecology of juvenile lake sturgeon in the St. Lawrence River system.

Transactions of the American Fisheries Society 135:1044-1055. 65

16. Pollock, M.S., M. Carr, N.M. Kreitals, and I.D. Phillips. 2014. Review of a

species in peril: what we do not know about lake sturgeon may kill them.

Environmental Reviews 23:30-43.

17. Stelzer, R.S., H.G. Drecktrah, M.P. Shupryt, and R.M. Bruch. 2008. Carbon

sources for lake sturgeon in Lake Winnebago, Wisconsin. Transactions of the

American Fisheries Society 137:1018-1028.

18. Vladykov, V.D. 1955. A comparison of Atlantic sea sturgeon with a new

subspecies from the Gulf of Mexico (Acipenser oxyrhynchus de sotoi). Journal of

the Fisheries Research Board of Canada 12:754-761.

19. Volkman, E.T., K.L. Pangle, D.A. Rajchel, and T.M. Sutton. 2004. Hatchery

performance attributes of juvenile lake sturgeon fed two natural food types. North

American Journal of Aquaculture 66:105-112.

66

Pallid Sturgeon, Scaphirhynchus albus (Forbes & Richardson, 1836)

Etymology and Synonyms: Scaphirhynchus = scaphe, Greek for “shovel”, and rhynchus, Greek for “snout”; albus = Latin for “white”, referring to the coloring of the species.

Description: Body fusiform, elongate, slightly arched and robust dorsally, and ventrally flattened. Dorsally grayish-blue to light brown; laterally tan; ventrally white; body with five rows of large scutes; dorsal scutes 14-18, lateral scutes 40-48, ventral scutes 9-13.

Head depressed. Snout large, elongate, flattened, and shovel-shaped. Eyes small, placed dorsolaterally; small spiracles absent slightly posterior to eyes. Mouth inferior with 4 fleshy lobes on lower lip. Four barbels present with accessory fringes; placed anterior to mouth on ventral side of head with two outer barbels posterior to two inner, thinner, and weakly fringed barbels. Gill rakers 13-14. Dorsal fin placed posteriorly on body with 37-

43 rays. Adipose fin absent. Caudal peduncle long and narrow; exposed portions of skin between scutes absent. Caudal fin heterocercal with upper lobe extending into a slight filament, often shortened in adults. Anal fin with 24-29 rays. Pelvic fin positioned posterior on body. Pectoral fins larger than pelvic or anal fins. Lateral line complete through upper lobe of caudal fin. Juveniles and adults similar in appearance.

Similar Species: Closely resembles and occurs sympatrically with the Shovelnose

Sturgeon. Shovelnose Sturgeon have barbels positioned in a straight, transverse line, instead of the inner barbels positioned anteriorly to outer barbels like on the Pallid

Sturgeon. Additionally, Shovelnose Sturgeon have scale-like scutes on the belly, and lower counts of rays in their dorsal fin (36 or fewer) and anal fin (23 or fewer); these counts slightly overlap with the counts of dorsal fin and anal fin rays in Pallid Sturgeon. 67

Lake Sturgeon barbels are not fringed, and the caudal peduncle is thicker with exposed areas of skin and spiracles not covered by bony plates.

Distribution and Habitat: Native to the Missouri and Mississippi Rivers and their major lower tributaries. Found in the mainstem Missouri River throughout North Dakota and

South Dakota. Roughly 51% of their range has been channelized, 28% has been impounded, and the remaining 21% occurs below dams which alter flow, sedimentation, and water temperature.8 Consequently, this species has declined dramatically and has been listed as federally endangered since 1990.15 Prefers areas with firm sand or silt substrate. Occur frequently in mid- and main-channels, areas around wing dikes, and occasionally islands with depths <1.5-4.0 m (4.92-13.12 ft) deep with strong current and highly turbid water.3,5,10 Lethal thermal maxima for age-0 individuals is related to body mass and acclimation temperature.5 Due to their limited swimming abilities early in life, young Pallid Sturgeon rely on low water velocities (<0.6 m/s) near the bottom generally in water depth <1.5 m (4.92 ft).1,16

Reproduction: Low abundance of juveniles in the Missouri River implies limited natural reproduction and recruitment of the species. Lack of recruitment may be related to scarce drift distance for larvae.16 Spawning takes place from late May to early June in the upper

Missouri River, and late April to early May in the lower Missouri and middle Mississippi

Rivers.9 Males reach sexual maturity at ages 5-7; females 9-20.9,14 Minimum length-at- maturity has been reported as 798 mm (31.4 in) FL for males, and 788 mm (31.02 in) FL for females.14 Spawning does not take place annually, and females may wait several years before spawning again.9,14 Detailed information on spawning habitat is scarce, but broadcast spawning is known to take place in shallow, rocky areas adjacent to the main 68 channel. Larvae drift in free-flowing water for 11-17 days before settling on the bottom substrate.11 Predation of age-0 Pallid Sturgeon, 30-45 mm (1.18-1.77 in) FL by Channel

Catfish and Smallmouth Bass has shown to be low, especially when alternative small prey is present, like Fathead Minnow.6 This leads to the assumption that predation is not limiting recruitment and survival of young.6

Age and Growth: Growth known to be rapid 13-48 days post-hatch, and then remain slow until the end of the first growing season (aprox. 100-110 days) when juveniles are

120-140 mm (4.72-5.51 in) FL.2 Length-at-age may vary depending on time of hatch, water temperature, and duration of the growing season.2 Juvenile optimal growth temperature is 25-28 °C (77-82.4 °F).4 Length-at-age for stocked individuals were reported as: age-1, 278 mm (10.94 in) FL; age-2, 315 mm (12.40 in) FL; age-3, 351 mm

(13.82 in) FL; age-4, 387 mm (15.24 in) FL; age-5, 424 mm (16.69 in) FL; age-6, 460 mm (18.11 in) FL.2,13 Capable of reaching 1,638 mm (64.48 in) FL.12 Long-lived species; individuals believed to reach over 50 years.

Food and Feeding: Benthic predator; uses barbels to detect prey. Wild feeding young- of-year primarily consume Diptera larvae, Diptera pupae, Ephemeroptera nymphs and

Chironomidae larvae.2,16,17 Juvenile optimal feeding temperature is 25-28 °C (77-82.4

°F).4 Individuals <600 mm (23.6 in) FL primarily consume macroinvertebrates, but undergo an ontogenetic diet shift by increasing their dependency on prey fishes once they exceed 600 mm (23.6 in) FL, or between ages 5 and 7.7

Literature Cited: 69

1. Adams, S.R., G.L. Adams, and G.R. Parsons. 2003. Critical swimming speed and

behavior of juvenile shovelnose sturgeon and pallid sturgeon. Transactions of the

American Fisheries Society 132:392-397.

2. Braaten, P.J., D.B. Fuller, R.D. Lott, T.M. Haddix, L.D. Holte, R.H. Wilson, M.L.

Barton, J.A. Kalie, P.W. DeHaan, W.R. Ardren, R.J. Holm, and M.E. Jaeger.

2012. Natural growth and diet of known-age pallid sturgeon (Scaphirhynchus

albus) early life stages in the upper Missouri River basin, Montana and North

Dakota. Journal of Applied Ichthyology 28:496-504.

3. Bramblett, R.G., and R.G. White. 2001. Habitat use and movements of Pallid and

Shovelnose Sturgeon in the Yellowstone and Missouri Rivers in Montana and

North Dakota. Transactions of the American Fisheries Society 130:1006-1025.

4. Chipps, S.R., R.A. Klumb, and E.B. Wright. 2008. Development and application

of juvenile pallid sturgeon bioenergetics model. State Wildlife Grant Program,

Study T-24_R Study No. 2424:40pp.

5. Deslauriers, D., L. Heironimus, and S.R. Chipps. 2016. Lethal thermal maxima

for age-0 Pallid and Shovelnose Sturgeon: Implications for shallow water habitat

restoration. River Research and Applications 32:1872-1878.

6. French, W.E., B.D.S. Graeb, S.R. Chipps, K.N. Bertrand, T.M. Selch, and R.A.

Klumb. 2010. Vulnerability of age-0 pallid sturgeon Scaphirhynchus albus to fish

predation. Journal of Applied Ichthyology 26:6-10.

7. Grohs, K.L., R.A. Klumb, S.R. Chipps, and G.A. Wanner. 2009. Ontogenetic

patterns in prey use by pallid sturgeon in the Missouri River, South Dakota and

Nebraska. Journal of Applied Ichthyology 25:48-53. 70

8. Keenlyne, K.D. 1989. A report on the pallid sturgeon. U.S. Fish and Wildlife

Service, Pierre, South Dakota.

9. Keenlyne, K.D., and L.G. Jenkins. 1993. Age at sexual maturity of the Pallid

Sturgeon. Transactions of the American Fisheries Society 122:393-396.

10. Koch, B., R.C. Brooks, A. Oliver, D. Herzog, J.E. Garvey, R. Hrabik, R.

Colombo, Q. Phelps, and T. Spier. 2012. Habitat selection and movement of

naturally occurring Pallid Sturgeon in the Mississippi River. Transactions of the

American Fisheries Society 141:112-120.

11. Kynard, B., E. Parker, D. Pugh, and T. Parker. 2007. Use of laboratory studies to

develop a dispersal model for Missouri River pallid sturgeon early life intervals.

Journal of Applied Ichthyology 23:365-374.

12. Shuman, D.A., D.W. Willis, and S.C. Krentz. 2006. Application of a length-

categorization system for Pallid Sturgeon (Scaphirhynchus albus). Journal of

Freshwater Ecology 21:71-76.

13. Shuman, D.A., R.A. Klumb, G.A. Wanner, R.H. Wilson, K.D. Steffensen, W.J.

Doyle, W.M. Gardner, T. Haddix, M. Ruggles, P.T. Horner, M.E. Jaeger, and S.

Stukel. 2011. Pallid sturgeon size structure, condition, and growth within the

Missouri River basin. Journal of Applied Ichthyology 27:269-281.

14. Steffensen, K.D., M.A. Pegg, and G.E. Mestl. 2013. Population characteristics of

pallid sturgeon (Scaphirhynchus albus (Forbes & Richardson, 1905)) in the

Lower Missouri River. Journal of Applied Ichthyology 29:687-695. 71

15. U.S. Fish and Wildlife Service. 1990. Endangered and threatened wildlife and

plants; determination of endangered status for the pallid sturgeon. U.S. Federal

Regulation 55:36641-36647.

16. USFW. 2003. Amendment to the 2000 Biological Opinion on the operation of the

Missouri River main stem reservoir system, operation and maintenance of the

Missouri River bank stabilization and navigation project, and operation of the

Kansas River reservoir system. Omaha District, NE, USA: U.S. Fish and Wildlife

Service.

17. Wanner, G.A., D.A. Shuman, and D.W. Willis. 2007. Food habits of juvenile

pallid sturgeon and adult shovelnose sturgeon in the Missouri River downstream

of Fort Randall Dam, South Dakota. Journal of Freshwater Ecology 22:81-92.

72

Shovelnose Sturgeon, Scaphirhynchus platorynchus (Rafinesque, 1820)

Etymology and Synonyms: Scaphirhynchus = scaphe, Greek for “shovel”, and rhynchus, Greek for “snout”; platorynchus = platy, Greek for “broad” or “flat”, and rhynchus, Greek for “snout”.

Description: Body fusiform, elongate, slightly dorsoventrally flattened. Dorsally light gray, light brown, brown or olive; laterally light brown to tan; ventrally white; body with five rows of large scutes; dorsal scutes 13-19, lateral scutes 38-50, ventral scutes 9-14.

Head depressed. Snout long, flattened and shovel-shaped. Eyes small, placed dorsolaterally on head; small spiracle slightly posterior to eyes absent. Mouth inferior, 4 fleshy lobes on lower lip. Barbels present, 4 with accessory fringes present anterior to mouth in a straight line; two inner barbels thick and strongly fringed. Frenum absent. Gill rakers 11-14. Dorsal fin placed posteriorly on body with 29-36 rays. Adipose fin absent.

Caudal peduncle long and narrow; exposed portions of skin between scutes absent.

Caudal fin heterocercal with upper lobe extending into a slight filament, often shortened in adults. Anal fin with 18-23 rays. Pelvic fins positioned posteriorly on body. Pectoral fins larger than pelvic or anal fins. Lateral line complete through upper lobe of caudal fin.

Spawning adults similar to non-spawners. Juveniles similar to adults.

Similar Species: Closely resembles and occurs sympatricly with the Pallid Sturgeon.

Pallid Sturgeon have two sets of barbels with accessory fringes present, with inner barbels positioned anteriorly to the outer barbels, and the two inner barbels thin and slightly fringed. Pallid Sturgeon also lack any scutes on the ventral side, have a dorsal fin with 37-43 rays, and an anal fin with 23-28 rays. Lake Sturgeon have a more short, rounded, and conical snout, smooth barbels, a short and thick caudal peduncle with 73 exposed portions of skin present between scutes, and the upper lobe of the caudal fin does not extend into a long filament.

Distribution and Habitat: Native to the upper Missouri and Mississippi River basins to the confluence of the Gulf of Mexico.11,13 Although Shovelnose Sturgeon are the most widespread sturgeon species in North America, their abundance has been declining due to anthropogenic factors such as pollution, habitat alteration, and overharvesting.11 Occur within the mainstem and direct tributaries of the Missouri River throughout North and

South Dakota. Inhabit the main channel areas of large rivers and their main tributaries with swift water over sand, gravel, and cobble substrate.4,9 Often found within pools located downstream of sandbars, dams, or wing dams.11 Juvenile habitat use is very similar to adults.9

Reproduction: The spawning season of Shovelnose Sturgeon is protracted, primarily occurring from late April to June, however the act may be extended into the fall or be bimodal, when fall conditions are similar to those during the spring.11,13 Spawning migrations consist of movements made upstream into smaller tributaries.1 Spawning is known to take place at water temperatures 16.9-20.5°C (62.42-68.9°F), and occur over hard substrates within tributaries of main rivers with increased levels of flow and depth.11

Experience delayed maturation, although Shovelnose Sturgeon mature earlier than other species of sturgeon due to their small size.13 Males reach sexual maturity at ages 5-8, and females at ages 7-9.11,13 Males generally spawn every other year, and females typically once every 3-4 years.13 Fecundity of females from the middle Mississippi River ranged

5,733-81,842 eggs, but averages 29,573 eggs and 21.7 eggs/g of body weight.13 No nest is 74 constructed and no parental care is given. Eggs adhesive, range in color from yellow, white, or black, and are roughly 2.27-2.5 mm (0.09-0.10 in) in diameter.1,5

Age and Growth: Smallest of the North American sturgeon species. Most Shovelnose

Sturgeon do not exceed 2.5 kg (5.51 lbs.), however individuals from the upper Missouri

River are reported to reach over 7 kg (15.43 lbs.).11 Shovelnose Sturgeon from the

Missouri River are characterized as long-lived and slow-growing, unlike individuals from the Mississippi River basin that tend to be young and experience faster growth.10 These differences in growth rates are likely related to regional environmental and anthropogenic factors.10 Mean lengths-at-age from the Missouri River are reported as: age-1, 213 mm

(8.39 in) TL; age-2, 274 mm (10.79 in) TL; age-5, 399 mm (15.71 in) TL; age-10, 503 mm (19.80 in) TL.6 Capable of reaching 1200 mm (47.24 in) TL.2 Estimated longevity is

43 years.7

Food and Feeding: Opportunistic benthivores. Forage by using their barbels to detect prey, and then capture prey by using their inferior mouth to suction feed. In the downstream segments of the Missouri River, Shovelnose Sturgeon do not experience ontogenetic shifts in the diet.8 Adults within the Missouri River mainly consume a variety of aquatic arthropods including Trichopetra, Diptera, and Ephemeroptera larvae, but also are known to consume Odonata, Coleoptera, and fish eggs.12,14 Seasonal shifts in the diet are likely influenced by rates and timing of discharge.12 Feeding behavior in the Missouri

River has been characterized by individuals restricted to benthic foraging in the late spring and summer, feeding on major components of the drift in the fall, and consuming a greater diversity of aquatic and terrestrial invertebrates in the winter months.12 Age-0 individuals feed primarily on Diptera and Ephemeroptera larvae.3 75

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Braaten, P. J., Fuller, D. B. and McClenning, N. D. (2007), Diet composition of

larval and young-of-year shovelnose sturgeon in the Upper Missouri River.

Journal of Applied Ichthyology, 23: 516–520.

4. Bramblett, R.G., and R.G. White. 2001. Habitat use and movements of pallid and

shovelnose sturgeon in the Yellowstone and Missouri rivers in Montana and

North Dakota. Transactions of the American Fisheries Society 130:1006-1025.

5. Bryan, J.L., M.L. Wildhaber, D.M. Papoulias, A.J. DeLonay, D.E. Tillitt, and

M.L. Annis. 2007. Estimation of gonad volume, fecundity, and reproductive stage

of shovelnose sturgeon using sonography and endoscopy with application to the

endangered pallid sturgeon. Journal of Applied Ichthyology 23:411-419.

6. Carlander, K.D. 1969. Handbook of freshwater fishery biology. Iowa State

University Press, Ames. 752pp.

7. Everett, S.R., D.L. Scarnecchia, G.J. Power, and C.J. Williams. 2003. Comparison

of age and growth of shovelnose sturgeon in the Missouri and Yellowstone rivers.

North American Journal of Fisheries Management 23:230-240.

8. French, W.E., B.D.S. Graeb, K.N. Bertrand, S.R. Chipps, and R.A. Klumb. 2013.

Size-dependent trophic patterns of pallid sturgeon and shovelnose sturgeon in a

large river system. Journal of Fish and Wildlife Management 4:41-52. 76

9. Gerrity, P.C., C.S. Guy, and W.M. Gardner. 2008. Habitat use of juvenile pallid

sturgeon and shovelnose sturgeon with implications for water-level management

in a downstream reservoir. North American Journal of Fisheries Management

28:832-843.

10. Hamel, M.J., J.J. Spurgeon, M.A. Pegg, J.J. Hammen, and M.L. Rugg. 2014a.

Hydrologic variability influences local probability of Pallid Sturgeon occurrence

in a Missouri River tributary. River Research and Applications 32:320-329.

11. Keenlyne, K.D. 1997. Life history and status of the shovelnose sturgeon,

Scaphirhynchus platorynchus. Environmental Biology of Fishes 48:291-298.

12. Modde, T., and J.C. Schmulbach. 1977. Food and feeding behavior of the

shovelnose sturgeon, Scaphirhynchus platorynchus, in the unchannelized

Missouri River, South Dakota. Transactions of the American Fisheries Society

106:602-608.

13. Tripp, S.J., Q.E. Phelps, R.E. Colombo, J.E. Garvey, B.M. Burr, D.P. Herzog, and

R.A. Hrabik. 2009. Maturation and reproduction of shovelnose sturgeon in the

middle Mississippi River. North American Journal of Fisheries Management

29:730-738.

14. Wanner, G.A., D.A. Shuman, and D.W. Willis. 2007. Food habits of juvenile

pallid sturgeon and adult shovelnose sturgeon in the Missouri River downstream

of Fort Randall Dam, South Dakota. Journal of Freshwater Ecology 22:81-92.

77

CHAPTER 5

FAMILY POLYODONTIDAE

Introduction

Only two extant species, both freshwater inhabitants, make up the ancient paddlefish family, Polyodontidae: the North American Paddlefish, Polyodon spathula, and the Chinese Paddlefish, Psephurus gladius. The Chinese Paddlefish is critically endangered and has not been reported in more than a decade, leading to the possibility that the species may have gone extinct. Originally, paddlefish were thought to be sharks when first discovered, given their scaleless body, cartilaginous skeleton and deeply forked heterocercal tail. However, paddlefish are a group of ancient bony fishes that belong in the order, Acipenseriformes, along with the closely related species of sturgeon from the family Acipenseridae. Paddlefish are also commonly called “spoonbill catfish” because of their spoon-like rostrum and their heterocercal caudal fin, similar to the caudal fin of a Channel Catfish, Ictalurus punctatus. However, this common name is rather misleading since the paddlefish family and catfish family, Siluridae, are not related. Both extant species of paddlefish have earned their common names in reference to their long and extended paddle or spatula-like rostrum, hence the North American Paddlefish’s specific epithet, spathula. Another defining feature the two members of the family share in addition to their rostrum, cartilaginous skeleton and heterocercal tail is the absence of the jawbones, or the maxillary and premaxillary bones.

The North American Paddlefish, native to the Missouri River in the Dakotas, has a native distribution that spreads across the Mississippi and Missouri River basins in the 78 east and central United States from New York to Montana, and south throughout the

Mobile basin to the Gulf of Mexico. The Chinese Paddlefish occurs within the Yangtze-

Kiang River drainage in southcentral and eastern China. Paddlefish inhabit large to medium sized turbid to semi-turbid rivers and reservoirs, often in areas with low water velocity below structures or sandbars with sand, gravel or silt substrates. Although North

American Paddlefish are still found across their native and vast distribution in the United

States, populations along the Gulf Coast and within the Mississippi River continue to be a concern. Over the past couple of decades, the North American Paddlefish has been negatively affected by anthropogenic habitat alterations including river channelization, dam construction and impoundments, which alter and affect migratory patterns.

Overharvesting has also been a contributing issue, with many individuals being taken for their roe.

The North American Paddlefish is planktivorous and forages by using a method called ram ventilation. The lack of jawbones requires and allows the paddlefish to have their mouths slightly open while continuously swimming, allowing phytoplankton and zooplankton to enter the mouth cavity and be filtered out. Excess water then passes over the gills as a form of ventilation. Paddlefish also have the ability to detect their prey using the numerous electrosensory receptors that cover the dorsal and ventral sides of the rostrum. Unlike the North American Paddlefish, the Chinese Paddlefish feeds on larger aquatic invertebrates and small fish. The North American Paddlefish migrates upstream to spawn in the springtime. Males generally spawn every year, unlike females who spawn every 2-3 years. Similar to sturgeon, paddlefish have delayed maturation, which is an important life history trait to take into account with how they are managed. In the 79

Dakotas, paddlefish are listed as a sportfish given their stable and self-sustaining populations. With the purchase of a tag, they can be sought after using a snagging method.

80

Paddlefish, Polyodon spathula (Walbaum, 1792)

Etymology and Synonyms: Poly = many, odon = tooth, in reference to the numerous gill rakers; spathula = blade, or paddle, in reference to the shape of the rostrum.

Description: Body thick, robust, and round in cross section. Dorsally and laterally blueish gray to black, often mottled; ventrally cream to white. Head broad. Opercle extends posteriorly past pectoral fins, and tapers into an elongate, pointed flap. Snout

(called a rostrum) long, paddle-shaped (roughly 1/3 of body length in adults), covered with small sensory pores. Eyes small. Spiracle present, posterior of eye. Mouth large, subterminal. Teeth small; present on upper and lower jaws, and on basal portions of gill arches in juveniles; adults toothless. Barbles present, not obvious; one small pair on ventral side of rostrum. Gill rakers numerous, long, and slender. Dorsal fin set posterior on body; insertion between pelvic an anal fins; no spines. Adipose fin absent. Caudal peduncle short, thick. Caudal fin heterocercal, strongly forked. Anal fin with concave distal end. Pelvic fin abdominal. Pectoral fins fleshy. Lateral line complete. Leather-like skin with scales absent, except for single patch of ganoid scales at base of upper lobe of caudal fin. Juveniles similar in appearance to adults with a rostrum greater than 1/3 body length.

Similar Species: No other species within the Dakotas have similar features. Paddlefish belong to the family Polyodontodae, which contains only one other species, the Chinese

Paddlefish (Psephurus gladius), which is native to the Yangtze-Kiang River.9

Distribution and Habitat: Native to the eastern and central United States in the

Mississippi-Missouri River basins and all their major tributaries as far south as the Gulf of Mexico. Populations still occur over much of the native range, although the species has 81 been negatively affected over time by river channelization, dam construction, dewatering, pollution, and overharvest for their roe.2,4,8 Occurs within large to medium-sized rivers, and reservoirs in open waters near side channels, areas below structures and sandbars, bays, and eddies with low current velocities below 0.3 m-sec.4,7,16 Habitats selected are known to vary seasonally and annually in response to discharge and temperature.4,12,16

Most often found over gravel, sand, and silt substrates, in turbid waters with depths averaging 3.0 m (9.8 ft).8,12

Reproduction: Upstream spawning migrations sometimes occurring over hundreds of miles to find suitable spawning areas begin when water temperatures near 10 °C (50 °F) and flow increases.8,15 Successful spawning is highly dependent on photoperiod, water temperature, and flow; if any of these factors are not satisfactory, females resorb their eggs.4,15 Males known to spawn each year; females every 2-3 years, which may be necessary to acquire the energy to produce the large egg masses.15 Sexual maturity known to vary with latitude, and is much longer than many other freshwater fishes.2 Males known to mature before females, often at ages 4-9, and females ages 6-12.2 Spawning takes place in the early spring in water temperatures 10-16 °C (50-61 °F) near the surface over gravel bars and hard bedrock that are free of silt.10,15 Information on spawning behavior is limited. Fertilized eggs are white, demersal, adhesive, and roughly 2.0-4.0 mm (0.08-0.16 in) in diameter.10,12 Fecundity is large, but considered variable by similarly-sized females. 11,15 Estimates of weight-specific fecundity from Arkansas range from 7,794-30,247 eggs/ kilogram (3,543-13,749 eggs/ lb.) of body weight.6 Hatching is directly related to water temperature.4 In water temperatures of 18-21 °C (64-70 °F), 82 hatching took place in 6-7 days.10 In water temperatures of 11-14 °C (52-57 °F), hatching took place in 12-14 days.4

Age and Growth: Newly hatched larvae average 8.5 mm (0.3 in) TL.5,10 Growth is rapid for the first five years, and known to be directly related to food abundance and length of growing season.15 Growth rates are believed to be higher in reservoirs than rivers.8Adults often measured from anterior region of eye to the fork in the caudal fin (eye-fork length,

EFL), to eliminate errors when faced with damaged rostrums or caudal fins.14 Females are known to be heavier and longer than males.12 Mean length for age-1 individuals from

Lewis and Clark Reservoir was 192.0 mm (7.6 in) EFL.14 The mean weight and mean length of Paddlefish in-between Yankton, South Dakota and Ponca State Park, Nebraska were 5,900 g (13 lbs.) and 730 mm (29 in).12 Capable of reaching lengths over 2.2 m (6 ft) and weighing up to 72 kg (158 lbs.).1,3,4 Average longevity, 5-8 years, however it is not uncommon for them to reach >20 years old.4,9

Food and Feeding: Adults are mainly planktivorous while larval individuals actively feed on larger zooplankton and insects until their gill rakers are developed, often at lengths of 120-150 mm (4.72-5.91 in) TL.13 Juveniles and adults are non-selective filter feeders on larger zooplankton.13 Occasionally consume small insects, insect larvae, and small fish.5,14 Prey is detected using electrosensory receptors on the rostrum.17

Literature Cited:

1. Allardyce, D.A. 1992. Endangered and threatened wildlife and plants: findings on

petition to list the paddlefish. Federal Register 57:43676-43682.

2. Carlson, D.M., and P.S. Bonislawsky. 1981. The paddlefish (Polyodon spathula)

fisheries of the Midwestern United States. Fisheries 6(2):17-27. 83

3. Epifanio, J.M., J.B. Koppelman, M.A. Nedbal, and D.P. Philipp. 1996.

Geographic variation of paddlefish allozymes and mitochondrial DNA.

Transactions of the American Fisheries Society 125(4):546-561.

4. Jennings, C.A., and S.J. Zigler. 2000. Ecology and biology of paddlefish in North

America: historical perspectives, management approaches, and research priorities.

Reviews in Fish Biology and Fisheries 10:167-181.

5. Jennings, C.A., and S.J. Zigler. 2009. Biology and life history of Paddlefish in

North America: an update. American Fisheries Society Symposium 66:1-22.

6. Leone, F.J., J.N. Stoeckel, and J.W. Quinn. 2012. Differences in paddlefish

populations among impoundments of the Arkansas River, Arkansas. North

American Journal of Fisheries Management 32:731-744.

7. Moen, C.T., D.L. Scarnecchia, and J.S. Ramsey. 1992. Paddlefish movements and

habitat use in Pool 13 of the upper Mississippi River during abnormally low river

stages and discharges. North American Journal of Fisheries Management, 12:744-

751.

8. Paukert, C.P., and W.L. Fisher. 2001. Characteristics of Paddlefish in a

Southwestern U.S. Reservoir, with Comparisons of Lentic and Lotic Populations.

Transactions of the American Fisheries Society, 130(40:634-643.

9. Pflieger, W.L. 1975. The Fishes of Missouri. Missouri Department of

Conservation, Jefferson City, Missouri, 343 pp.

10. Purkett, C.A., Jr. 1961. Reproduction and early development of the paddlefish.

Transactions of the American Fisheries Society, 90:125-129. 84

11. Reed, B.C., W.W. Kelso, and D.A. Rutherford. 1992. Growth, fecundity, and

mortality of paddlefish in Louisiana. Transactions of the American Fisheries

Society, 121:374-384.

12. Rosen, R.A. 1976. Distribution, age and growth, feeding ecology of paddlefish

(Polyodon spathula) in unaltered Missouri River, South Dakota. MSc Thesis.

South Dakota State University, Brookings, South Dakota, 95pp.

13. Rosen, R.A. and Hales, D.C. 1981. Feeding of paddlefish, Polyodon spathula.

Copeia 1981(2):441-455.

14. Ruelle, R., and P. Hudson. 1977. Paddlefish (Polyodon spathula): growth and

food of young of the year and a suggested technique for measuring length.

Transactions of the American Fisheries Society 106:609-613.

15. Russell, T.R. 1986. Biology and life history of the paddlefish-a review. In:

Dillard, J.G., Graham, L.K. and Russell, T.R. (eds.), The Paddlefish: Status,

Management and Propagation. American Fisheries Society, North Central

Division, Special Publication 7, Bethesda, Maryland, pp.2-21.

16. Southhall, P.D., W.A. Hubert. 1984. Habitat use by adult paddlefish in the upper

Mississippi River. Transactions of the American Fisheries Society, 113:125-131.

17. Wilkins, L.A., and M.H. Hofmann. 2007. The paddlefish rostrum as an

electrosensory organ: a novel adaptation for plankton feeding. Bioscience 57:399-

407.

85

CHAPTER 6

FAMILY LEPISOSTEIDAE

Introduction

The Gar family, Lepisosteidae, consists of seven extant species within the two genera, Lepisosteus and Atractosteus. Gar are an ancient group of fishes, with fossils from the Cretaceous period dating back to roughly 75-100 million years ago from western North America, Europe, Africa, Asia and South America. The distribution of Gar nowadays is much more limited, with only 5 species occurring in North America, one species in Central America, and one species in Cuba. The majority of the seven extant species of Gar are freshwater inhabitants; however, some species like the Alligator Gar,

Atractosteus spatula, is occasionally found within brackish or marine waters along the

Gulf of Mexico. Two species, the Longnose Gar, Lepisosteus osseus, and the Shortnose

Gar, Lepisosteus platostomus, occur in the Dakotas.

Gars have a rather distinct general body plan compared to other families of fishes in the Dakotas. They have elongate, cylindrical, and torpedo-like bodies that are covered in hard, rhomboid-shaped ganoid scales, a dorsal and anal fin that is set far posterior on the body, and an abbreviate-heterocercal caudal fin. The snouts of Gar are also elongate and are lined with multiple sharp teeth along both jaws that help to quickly devour their prey. The swim bladder of Gar is lung-like, which allows Gar to gulp air. This action paired with the normal method of breathing through the gills makes them bimodal. The amount of air that Gar gulp often depends on the oxygen availability of their habitat and water temperature, with more facultative aerial breathing occurring in low oxygenated 86 waters with higher temperatures. This breathing adaptation is likely the main reason gars have survived fluctuating climactic stages throughout their existence.

When foraging, Gar may stalk their prey for several minutes before they exhibit a sit-and-wait behavior and then ambush their prey. When ambushing, the body forms a rough “S” shape, with the head turning quickly and the jaws wide open to grab the prey.

Once the Gar have the prey in their mouth, their heads shake it back and forth impairing the prey with their sharp teeth. All species of Gar are predators, with the majority of adults being piscivorous on smaller fish, although some species are also known to consume macroinvertebrates. Gar inhabit quiet, low current to stagnant areas within large rivers, streams, reservoirs, lakes and ponds with coarse substrate. During the spawning season, which occurs during the late spring and early summer, Gar migrate to shallower areas to spawn over coarse substrate or beds of vegetation. No nest is constructed, but the adhesive and demersal eggs adhere to vegetation or other debris while they develop. The eggs also have a gelatinous coating, which are known to be poisionous or toxic to other vertebrates, including humans.

The Longnose Gar are known to be absent from North Dakota, and are rather uncommon in South Dakota as their populations seem to be declining. Shortnose Gar occur throughout the Missouri and James Rivers in both North and South Dakota, although they are more abundant in southeastern South Dakota. Although they are a less popular sport fish in the Dakotas, their toothy mouths make them harder to hook, presenting a challenge to anglers.

87

Longnose Gar, Lepsisosteus osseus (Linnaeus, 1758)

Etymology and Synonyms: Lepisosteus = scales of bone; osseus = bony

Description: Body fusiform, elongate, slender, cylindrical. Coloration varies greatly with habitat; dorsally olive to dark green; laterally olive, brown, or tan with few small dark spots posteriorly; ventrally cream to white; large, dark spots present on dorsal, anal, and caudal fins; small spots occasionally on ventral side of jaw. Head elongate. Snout long, slender, dorso-ventrally flattened; width at nostrils less than diameter of eye; narrowest width less than 1/10th the distance from tip of snout to eye. Eye large; directly posterior to opening of mouth. Mouth terminal to subterminal. Teeth small, numerous, villiform; present on both jaws. Barbels absent. Gill rakers 20-31, rudimentary. Dorsal fin set far posterior. Adipose fin absent. Caudal peduncle short, slender. Caudal fin heterocercal with rounded distal end. Anal fin set far posterior; insertion anterior to anterior end of dorsal fin. Pelvic fin abdominal. Pectoral fin insertion directly posterior to opercle.

Lateral line scales 57-65; 47-55 predorsal scales; 17-24 transverse scales. Scales thick, ganoid (rhomboid). Air bladders connected to digestive tracts, allow for gulping of supplemental air from the surface. Juveniles similar to adults, but with distinct brown to black midlateral streak with white underlining from snout to base of caudal fin.

Similar Species: Closely resembles the Shortnose Gar. Shortnose Gar have a shorter, blunter, and wider snout; narrowest width greater than 1/10th the distance from tip of snout to eye. Often lighter in color than Longnose Gar, and have 20-23 transverse scales.

Distribution and Habitat: Occur throughout much of the eastern half of the United

States. Native to the southeastern edge of the Missouri River basin, the Mississippi River basin, and east to the Ohio and South-Atlantic Gulf basins. More common in fresh and 88 brackish water, but have been seen in saltwater with salinities up to 31ppt.9,11 Absent from North Dakota. Rather uncommon species in South Dakota; population has declined most likely due to increased levels of turbidity and siltation from construction of impoundements.6 Inhabit quiet, low-current areas of large rivers and streams, reservoirs, and lakes. Adults often caught or observed in deep open-water with moderate amounts of aquatic vegetation and structure over gravel and sand substrate. Young remain among vegetation during their first summer before moving to deeper waters as they mature.5

Reproduction: Spawning migrations take place April-July and have been called broad and extensive.7,9 Migrations to desired spawning grounds are known to be positively correlated with stream flow and water level, and negatively correlated with water temperature.7 In lakes and reservoirs, Longnose Gar migrate to shallower areas and smaller tributary streams to spawn.5,8 Individuals in rivers and streams often migrate upstream to smaller, higher-gradient streams.10,15 Spawning activity known to take place over deep, rocky stream reaches, or shallower areas with gravel bars in a peak water temperature of 19.5-21°C (67-70°F).1,10,15 Sexual maturity is reached at ages 2-4 in males, and ages 6-7 for females.4,5,7 Males known to occupy spawning grounds longer than females. One to several young males escort and nudge a single older female with their snouts to the spawning ground, where clutches of eggs are released and fertilized.5,7

No nest is prepared. Fecundity related to total body weight and total ovary weight; estimates range from 10,924-34,412 eggs per female.7 Eggs 2.50-3.20mm (0.09-0.13in) in diameter, greenish to gray, demersal and adhesive.1,4 Hatching takes place 7-10 days after spawning, but dependent on water temperature.5,12

89

Age and Growth: Newly hatched larvae 9-10mm (0.35-0.39in) TL.15 Growth rapid in the first year, reaching an approximate length of 400mm (in) at age-1.4,9,13 Growth significantly slower in both sexes after sexual maturity is reached.4,5,9 Females known to grow larger and live longer than males. Capable of reaching maximum lengths of roughly

1829cm (6ft). Average longevity 11 years for males; 22 years for females.3

Food and Feeding: Large apex, primarily piscivorous, ambush predators. Known to stalk victim before quickly striking to catch the prey between its jaws, followed by few thrashing movements. Digestion rates known to be slow.9 Feeding often takes place at night and near the surface. Juveniles feed primarily on small crustaceans, aquatic insects, and insect larvae until they reach a total length >11mm (4.6 in), when they become primarily piscivorous.2 Larger individuals primarily piscivorous and tend to prey on larger forage species such as shads, cyprinids, and to a lesser extent, sport fishes.9,14

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Echelle, A.A. 1968. Food habits of young-of-year longnose gar in Lake

Texacoma, Oklahoma. Southwestern Naturalist 13:45-50.

3. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

4. Ferrara, A.M. 2001. Life-history strategy of Lepisosteidae: implications for the

conservation and management of alligator gar. PhD dissertation, Auburn

University, Auburn, Alabama. 90

5. Hasse, B.L. 1969. An ecological life history of the longnose gar, Lepisosteus

osseus (Linnaeus), in Lake Mendota and several other lakes of southern

Wisconsin. PhD dissertation, University of Wisconsin, Madison.

6. Hoagstrom, C.W., C.A. Hayer, J.G. Kral, S.S. Wall, and C.R. Berry Jr. 2006. Rare

and declining fishes of South Dakota: a river drainage scale perspective.

Proceedings of the South Dakota Academy of Science 85:171-211.

7. Johnson, B.L., and D.B. Noltie. 1996. Migratory dynamics of stream-spawning

longnose gar (Lepisosteus osseus). Ecology of Freshwater Fish 5:97-107.

8. Kelley, S.W. 2012. Age and growth of spawning longnose gar (Lepisosteus

osseus) in a north central Texas reservoir. Western North American Naturalist

72:69-77.

9. McGrath, P.E. 2010. The life history of Longnose Gar, Lepisosteus osseus, an

apex predator in the tidal waters of Virginia. Doctoral dissertation, College of

William and Mary, Williamsburg, Virginia.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri

Department of Conservation, Jefferson City.

11. Schwartz, J. 2003. Longnose gar, Lepisosteus osseus (Family Lepisosteidae) in

North Carolina, especially the Cape Fear River. Journal of the North Carolina

Academy of Science 119:26-32.

12. Simon, T.P., and R. Wallus. 1989. Contributions to the early life histories of gar

(Actinopterygii: Lepidostidae) in the Ohio and Tennessee River basins with

emphasis on larval development. Transactions of the Kentucky Academy of

Science 50:59-74. 91

13. Sutton, T.M., A.C. Grier, L.D. Frankland. 2009. Stock structure and dynamics of

the Longnose gar and Shortnose gar in the Wabash River, Indiana-Illinois. Journal

of Freshwater Ecology 24:657-666.

14. Tyler, J.D., J.R. Webb, T.R. Wright, J.D. hargett, K.J. Mask, and D.R. Schucker.

1994. Food habits, sex ratios, and size of longnose gar in southwestern Oklahoma.

Proceedings of the Oklahoma Academy of Science 74:41-42.

15. Yeager, B.L., and R.T. Bryant. 1983. Larvae of the longnose gar, Lepisosteus

osseus, from the Little River in Tennessee. Journal of the Tennessee Academy of

Science 58:20-22.

92

Shortnose Gar, Lepisosteus platostomus (Rafinesque, 1820)

Etymology and Synonyms: Lepisosteus = Greek, lepis, meaning “scale”, and osteon, meaning “bone”, “scales of bone”; platystomus = Greek, platy, meaning “flat”, and stomus, meaning “mouth”.

Description: Body fusiform, elongate, slender, cylindrical. Dorsally olive green; laterally olive-silver to tan, small spots faint to absent posteriorly; ventrally cream to white. Paired fin spots typically absent and a few large dark spots present on dorsal, caudal, and anal fins. Head elongate. Snout dorso-ventrally flattened, short and broad; least snout width contained 10 or fewer times in the snout length. Eye large; placed laterally on head and directly posterior to opening of mouth. Mouth terminal to subterminal. Barbels absent.

Teeth are small, numerous, and villiform; present on both jaws. Gill rakers 16-25, rudimentary. Dorsal fin set far posterior. Adipose fin absent. Caudal peduncle short, slender. Caudal fin hemihomocercal with rounded distal end. Anal fin set far posterior on body; insertion slightly anterior to insertion of dorsal fin. Pelvic fins abdominal. Pectoral fin insertions directly posterior to opercle. Lateral line with 55-64 thick ganoid

(rhomboid) scales; 45-54 predorsal scales; 20-23 transverse scales. Swim bladder is connected to the digestive tract, allowing gulping of supplemental air from the surface.

Juveniles similar to adults, but have a darker mid-dorsal streak and more pronounced spots on caudal fin.

Similar Species: Shortnose Gar closely resemble Longnose Gar. Longnose Gar have a longer and more slender snout, with the least snout width contained 13 or more times in the snout length. Longnose gar also have small dark spots present posteriorly on lateral sides, and may have small spots present on ventral side of jaw. 93

Distribution and Habitat: Native to the Missouri River basin from eastern Montana, east into the Mississippi River basin to western Ohio, and south to the Gulf of Mexico.

Within the Dakotas, Shortnose Gar occur within the Missouri, James, Cannonball, Big

Sioux, and Vermillion river basins, and are more common in southeastern South Dakota.

Shortnose Gar inhabit quiet, low-current areas such as pools and backwaters in rivers, streams, and oxbows with sand and silt substrates. In lakes, Shortnose Gar often inhabit areas over sand bars and shoals.1 The species also tends to avoid areas with increased flow and heavy amounts of rooted aquatic vegetation.5 Shortnose Gar are more tolerant of siltation and higher turbidity than other gar species.5,8 Activity levels are increased at night when they migrate into shallower waters.2 Aerial breathing rates are known to increase with the size of the individual, throughout the night, and in warmer water temperatures greater than 15 °C (60 °F).12

Reproduction: Recent literature on reproduction habits and behavior is scarce. Shortnose

Gar spawning migrations are not known to consist of great distances, however large schools of individuals form just prior to spawning activity.3 During 1956 in South

Dakota, spawning occurred from late May to mid-June in water temperatures of 19-24 °C

(66-75 °F); in 1957 and 1958, spawning took place from late June to early July inferring that spawning activity is driven by water temperature.1,2 Sexual maturity is reached at age

3-4.1,8,14 Spawning takes place in shallow waters with no flow, often over beds of aquatic vegetation with no previous preparation of a nesting structure.5 Data on fecundity of

Shortnose Gar from the Dakotas, as well as the Midwest region is scarce to none.

Fecundity is known to increase with the length of the female.6 Eggs yellow green in 94 color, roughly 2.5 mm (0.1 in) in diameter, and released in small gelatinous and adhesive masses.9 Hatching takes place 8-9 days after spawning.8

Age and Growth: Newly hatched larvae roughly 8 mm (0.3 in) TL.4 Growth is rapid during the first year. Age at length from Lewis and Clark Lake, South Dakota was recorded as: age-1, 417 mm (16.4 in) TL; age-2, 486 mm (19.1 in) TL; age-3, 536 mm

(21.2 in) TL; age-4, 587 mm (23.1 in) TL; age-5, 605 mm (23.8 in) TL; age-6, 671 mm

(26.4 in) TL; age-7, 734 mm (28.9 in) TL.1,16 Capable of reaching lengths of 750-900 mm

(29.5-35.4 in) TL.15 Shortnose Gar are a more robust species than Longnose Gar, however average length is much shorter.1 Longevity up to 12 years.14

Food and Feeding: Primarily piscivorous, opportunistic, ambush predators. Prey and feeding habits are more diverse than other species of gar.11 Larval feeding takes place roughly 16 days post hatching.7,10 Juveniles feed near the surface on insect larvae, small crustaceans, and small fish.1,7,8 Adults mainly consume other fish species such as sunfish

(Lepomis spp.), and small Common Carp throughout the water column.7,13 Adults also ingest greater amounts of crayfish and insects such as cicadas, mayflies, and midges, which also play an important role in the diet composition.1,5,15 Dominance hierarchy and territorial defense while feeding has also been reported.15

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Carlander, K.D. 1969. Handbook of freshwater fishery biology, volume 1. Iowa

State University Press, Ames.

3. Coker, R.E. 1930. Studies of common fishes of the Mississippi River at Keokuk.

Bulletin of the US Bureau of Fisheries 1072:141-225. 95

4. Echelle, A.A., and C.D. Riggs. 1972. Aspects of the early life history of gars

(Lepisosteus) in Lake Texoma. Transactions of the American Fisheries Society

101:106-112.

5. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

6. Holloway, A.D. 1954. Notes on the life history and management of the Shortnose

and Longnose Gar in Florida waters. The Journal of Wildlife Management

18:438-449.

7. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

8. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

9. Potter, G.E. 1926. Ecological studies of the short-nose gar-pike (Lepisosteus

platostomus). University of Iowa Studies in Natural History 11:17-27.

10. Richardson, R.E. 1913. Observations on the breeding habitats of fishes at Havana,

Illinois, 1910 and 1911. Bulletin of the Illinois State Laboratory of Natural

History 11:117-127.

11. Robinson, H.W., and T.M. Buchanan. 1988. Fishes of Arkansas. University of

Arkansas Press, Fayetteville, 536 p.

12. Saksena, V.P. 1975b. Effects of temperature and light on aerial breathing of

shortnose gar, Lepisosteus platostomus. Ohio Journal of Science. 96

13. Shields, J.T. 1957. Report of fisheries investigations during the second year of

impoundment of Gavins Point Reservoir, South Dakota, 1956. South Dakota

game Fish and Parks, Dingell-Johnson Project F-1-R-6.

14. Sutton, T.M., A.C. Grier, and L.D. Frankland. 2009. Stock structure and dynamics

of longnose gar and shortnose gar in the Wabash River, Indiana-Illinois.

Freshwater Ecology 24:657-666.

15. Vokoun, J.C. 2000. Shortnose gar (Lepisosteus platostomus) foraging on

periodical cicadas (Magicicada spp.): territorial defense of profitable pool

positions. American Midland Naturalist 143:261-265.

16. Walburg, C.H. 1964. Fish population studies, Lewis and Clark Lake, Missouri

River, 1956-1962. US Fish and Wildlife Service Special Scientific Report,

Fisheries Number 482.

97

CHAPTER 7

FAMILY HIODONTIDAE

Introduction

The Mooneye family, Hiodontidae, comprises only two species, both of which occur in the Dakotas, the Mooneye, Hiodon tergisus, and the Goldeye, Hiodon alosoides.

The family Hiodontidae is only from North America, and although only two species are extant, the family is represented by several other fossilized species. At a first glance,

Hiodontids may initially resemble or be confused with members of the family Clupeidae, which comprises herrings and shads which also occur in the Dakotas. Members of both families are rather simple in their appearance and have silver bodies, a triangular shaped flap called an axillary process near the base of the pelvic fins, and transparent partial membranes that cover the anterior and posterior corners of the eye called the adipose eyelids. Hiodontids can easily be distinguished from Clupeids by the presence of a lateral line, which Clupeids lack, and the lack of pointed scales or serrated keel along the ventral side of the body, which Clupeids possess. The dorsal fin insertion on Hiodontids also aligns or nearly aligns with the insertion of the anal fin, unlike Clupeids where the dorsal fin insertion is distinctly positioned anterior to the insertion of the anal fin. Hiodontids also have small but noticeable sharp teeth present on both sets of jaws, as well as on the tongue and the roof of the mouth.

Both species of the Hiodontid family inhabit large rivers, tributaries, reservoirs and lakes. Mooneye are known to prefer more clear and quiet waters, whereas Goldeye are more tolerant of turbid and muddy backwaters with little to strong current. This may 98 explain the reason why Mooneye are far less frequent in South Dakota and are most often found throughout the Red River of the North in North Dakota. Spawning takes place during spring, when adults begin to migrate from deeper waters to more shallow and quiet areas of rivers, streams, and lakes. Due to the lack of an oviduct, the eggs are released inside the body cavity of females prior to spawning before they are released outside of the females body. No parental care is given, and the non-adhesive eggs and larvae develop as they drift downstream with the current. Both species feed opportunistically along the surface at night on small aquatic invertebrates such as zooplankton, and occasionally small fish.

Although they are not often targeted by anglers in the Dakotas, both Goldeye and

Mooneye are sometimes sought after, especially in Canada, as a delicacy. They also make great bait fish when fishing for catfish.

99

Goldeye, Hiodon alosoides (Rafinesque, 1819)

Etymology and Synonyms: Hiodon = Greek for “toothed tongue”, referring to the small teeth present on tongue; alosoides = alosa-, Latin for “shad”, -eides, Greek for

“resemblance”, referring to the species resemblance to the genus Alosa, that includes herrings and shads.

Description: Body strongly laterally compressed, elongate. Dorsally pale blue-green; laterally and ventrally solid silver to light silver-gold; fins without markings. Head fairly short. Snout blunt. Eye relatively large with yellow to gold iris; adipose eyelid present.

Mouth large, terminal, and oblique; maxillary extends below posterior margin of eye.

Teeth small, sharp; present on both jaws, tongue, and roof of mouth. Gill rakers short, thick; 15-17. Dorsal fin located posteriorly on body with 9-10 rays. Adipose fin absent.

Caudal peduncle short, thick. Caudal fin forked. Anal fin elongate, insertion slightly anterior to insertion of dorsal fin; 29-34 rays. Pelvic fin abdominal, insertion far anterior of dorsal fin insertion. Axillary process present, flap-like. Pectoral fins with 11-12 rays, no spines. Keel on ventral side lacking scales, not serrate; extends from anus to pectoral fins. Lateral line complete with 57-62 cycloid scales in series. Sexual dimorphism present in adults; anal fin strongly sickle-shaped in males, slightly bowl-shaped in females.

Similar Species: Closely resembles the Mooneye. Mooneye display 11-12 rays in the dorsal fin, with the dorsal fin insertion distinctly anterior of anal fin insertion. Mooneye also display an iris more silver in color, a maxillary extending to middle of the eye, and a ventral keel extending from the anus to the pelvic fins. Herring, Gizzard Shad, and

Alewife have serrated ventral keels, and lack teeth on the jaws and tongue. 100

Distribution and Habitat: Endemic and widely distributed across North America.

Native to the Hudson Bay drainage in the north, south to the Mississippi and Missouri

River basins, east to the Ohio River basin, and south to Louisiana and northern Texas.

Occurs throughout the Missouri River and its major tributaries in North Dakota and

South Dakota, as well as the Red River of the North basin in North Dakota. Inhabits large rivers, reservoirs, tributaries, and quiet backwaters with turbid, muddy water and little to strong current.4 Found most frequently just below the surface to midwater depths.

Overwinter in deeper waters of reservoirs, lakes, and rivers.7

Reproduction: Spawning migrations take place early spring; individuals move from deep to shallow areas and pools of lakes or streams, a few hundred meters from shoreline.1,4,7 Sexual maturity of males generally reached at age 2-3; females typically a year later.4 Spawning takes place annually and usually lasts 3-6 weeks.4 Spawning activity begins mid-May to early June, and has been observed very little since it is assumed to take place at night in highly turbid water, when water temperatures near 11°C

(51.8°F).7 Eggs are released within the body cavity of the female prior to spawning due to the lack of an oviduct.4 Fecundity from the upper Missouri River was reported as 4,288-

10,164 eggs/ female.3 Fertilized eggs semi-buoyant, non-adhesive, and roughly 2-4 mm

(0.08-0.16 in) in diameter. After hatching, larvae drift downstream with the current, and wind or turbulence may increase mortality.1,2

Age and Growth: Growth varies with location and length of growing season.4 Juveniles of both sexes grow at the same rate, but adult females grow faster than adult males.4

Warmer weather in spring and summer known to increase growth and development of young, in turn increasing survival rates.1 Mean length-at-age taken for Missouri River 101 backwaters in North Dakota are reported as: age-1, 121 mm (4.76 in) TL; age-2, 189 mm

(7.44 in) TL; age-3, 234 mm (9.21 in) TL; age-4, 276 mm (10.86 in) TL; age-5, 298 mm

(11.73 in) TL; age-6, 305 mm (12.00 in) TL; age-7, 314 mm (12.36 in) TL.5 Females generally live longer and attain greater total lengths than males. Adults average 381-432 mm (15-17 in) TL. Capable of reaching 508 mm (20 in) TL.8 Individuals from South

Dakota rarely exceed 907.2 g (2 lbs.).6 Longevity, 7-9 years; may be longer in more northern populations.3,4

Food and Feeding: Surface feeder. Mainly feeds after dark. Larvae primarily consume daphnia and copepods. Juveniles consume daphnia, copepods, corixids, as well as aerial insects, but variation likely depends on availability of food.2,4 Adults feed on zooplankton, a variety of aquatic and terrestrial insects, and occasionally small fishes.2

Selection of prey may vary depending on month and location.4

Literature Cited:

1. Donald, D.B. 1997. Relationship between year-class strength for Goldeyes and

selected environmental variables during the first year of life. Transaction of the

American Fisheries Society 126:361-368.

2. Donald, D.B., and A.H. Kooyman. 1977. Food, feeding habits, and growth of

Goldeye, Hiodon alosoides (Rafinesque), in waters of the Peace-Athabasca Delta.

Canadian Journal of Zoology 55:1038-1047.

3. Hill, W.J. 1966. Observations on the life history and movement of the Goldeye,

Hiodon alosoides, in Montana. M.S. Thesis, Montana State University, Bozeman.

4. Kennedy, W.A., and W.M. Sprules. 1967. Goldeye in Canada. Fisheries Research

Board of Canada Bulletin 161. 102

5. Moon, D.N., S.J. Fisher, and D.W. Willis. 1998. Goldeye recruitment and growth

on two Missouri River backwaters. Proceedings from the South Dakota Academy

of Science 77:139-144.

6. Neumann, R.M., and D.W. Willis. 1994. Guide to the Common Fishes of South

Dakota. South Dakota Department of game, Fish and Parks, Pierre, South Dakota

7. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

8. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

103

Mooneye, Hiodon tergisus (Lesueur, 1818)

Etymology and Synonyms: Hiodon = Greek for “toothed tongue”, referring to the small teeth present on tongue; tergisus = from the Greek word tergisus, meaning “polished”.

Description: Body strongly laterally compressed, elongate. Dorsally pale blue-green; lateral, and ventral sides solid silver to light silver-gold in color; fins without markings.

Head fairly short. Snout blunt. Eye relatively large with iris mainly silver in color. Mouth large, terminal and oblique; maxillary extends to the middle of the eye. Teeth small, sharp; present on both jaws, tongue, and roof of mouth. Gill rakers short, thick, knob- like; 15-17. Dorsal fin located posterior on body with 11-12 rays, no spines. Adipose fin absent. Caudal peduncle short, thick. Caudal fin forked. Anal fin elongate, insertion distinctly posterior to insertion of dorsal fin; 26-29 rays; enlarged anterior lobe present on male anal fin. Pelvic fin abdominal, insertion anterior of dorsal fin insertion. Axillary process present, flap-like. Pectoral fins with 13-15 rays, no spines. Keel on ventral side lacking scales, not serrate; extends from anus to pelvic fins. Lateral line complete with

52-57 cycloid scales. Sexual dimorphism present in adults; anal fin strongly sickle- shaped in males; slightly bowl shaped in females.

Similar Species: Closely resembles the Goldeye. Goldeye display 9-10 rays in the dorsal fin, with insertion slightly posterior to anal fin insertion. Goldeye also display an iris more yellow to gold in color, a maxillary extending below the posterior margin of the eye, and a ventral keel extending from the anus to the pectoral fins. Herring, Gizzard

Shad, and Alewife have serrated ventral keels, and lack teeth on the jaws and tongue.

Distribution and Habitat: Native to the Great Lakes and Hudson drainages from

Quebec to Alberta, as well as the Mississippi and Missouri Rivers and their major 104 tributaries south to the Gulf of Mexico. Occurs in the Dakotas, but much less frequent than Goldeye. Found within the Red River of the North drainage in North Dakota, as well as the Cannonball, Knife, and Little Missouri River drainages.11 Rarely found in South

Dakota, but has been documented from the Grand, Lower Big Sioux, and Vermillion

River drainages. Mooneye range and abundance has been negatively impacted due to increased sedimentation and fluctuations of flow from dams, making the species less frequent since they are less tolerant of silt and turbidity.4,8 Little information is available on habitat use. Young-of-year in rivers inhabit backwater pools with low velocity along riverbanks.6 Adults inhabit tributaries and reservoirs with clear, quiet water, and thrive in large clear rivers and near the tailwaters of locks and dams.9

Reproduction: Spawning migrations consist of numerous adults moving into quiet, clear streams overnight.1 Sexual maturity is reached between ages 3-5, with males maturing a year earlier than females.7 Spawning takes place annually, starting in spring and extending into mid-summer; southern populations spawn earlier than northern populations.12 Little information on spawning activity has been recorded, but it is known to take place in water temperatures 10-13°C (50-55°F), in clear, flowing water over coarse substrate.2,7,8 Mean fecundity typically 5,000-9,000 eggs/ female.7,8 Fecundity increases with size and age of female, and depends on a variety of physiological and environmental factors.7,8 Eggs are released within the body cavity of the female prior to spawning due to the lack of an oviduct.3 Complete spawners, all eggs released at one time.8 Eggs buoyant, non-adhesive, roughly 1.98-2.77 mm (0.08-0.11 in) in diameter, and develop as they drift downcurrent.2,7,8 105

Age and Growth: Slightly <7.0 mm (0.28 in) at time of hatching.10 Growth rapid during first summer, and known to be faster in the northern parts of the species range.6,8 Young- of-year may decrease more than 25% in body weight, and experience slower or stunted growth during the winter.6 Mean lengths-at-age taken from the lower Tallapoosa River in

Alabama were reported as: age-1, 121 mm (4.76 in) TL; age-2, 170 mm (6.69 in) TL; age-3, 206 mm (8.11 in) TL; age-4, 233 mm (9.17 in) TL; age-5, 254 mm (10.00 in) TL; age-6, 269 mm (10.59 in) TL; age-7, 281 mm (11.06 in) TL; age-8, 290 mm (11.41 in)

TL.8 Females generally live longer than males.7 Longevity 7-9 years.

Food and Feeding: Adults opportunistic, surface feeders. Feeding occurs throughout the year, although majority takes place during spring and autumn, and very little occurs during winter.5,6,7 Individuals over age-1 primarily consume insects including Corixids,

Coleopterans, Ephemeropterans, Dipterans, and Odonatans.5 Adults also known to consume zooplankton, crayfish, plant material, and occasionally small fishes.5 Diets of young similar to adults, however contain more zooplankton and immature insects indicating that initial feeding takes place further below the surface.6 Feeding takes place at night, although to a lesser extent than Goldeye.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville. 106

4. Freeman, M.C., Z.H. Bowen, K.D. Bovee, and E.R. Irwin. 2001. Flow and habitat

effects on juvenile fish abundance in natural and altered flow regimes. Ecological

Applications 11(1):179-190.

5. Glenn, C.L. 1975. Seasonal diets of Mooneye, Hiodon tergisus, in the Assiniboine

River. Canadian Journal of Zoology 53:232-237.

6. Glenn, C.L. 1978. Seasonal growth and diets of young-of-the-year Mooneye

(Hiodon tergisus) from the Assiniboine River, Manitoba. Transactions of the

American Fisheries Society 107(4)587-589.

7. Glenn, C.L., and R.R.G. Williams. 1976. Fecundity of Mooneye, Hiodon tergisus,

in the Assiniboine River. Canadian Journal of Zoology 54:156-161.

8. Katechis, C.T., P.C. Sakaris, and E.R. Irwin. 2007. Population demographics of

Hiodon tergisus (Mooneye) in the Lower Tallapoosa River. Southeastern

Naturalist 6(3):461-470.

9. Mettee, M.F., P.E. O’Neil, and J.M. Pierson. 1996. Fishes of Alabama and the

Mobile Basin. Geological Survey of Alabama Monograph 15, Tuscaloosa.

10. Snyder, D.E., and S.C. Douglas. 1978. Description and identification of Mooneye,

Hiodon tergisus, protolarvae. Transactions of the American Fisheries Society

107(4):590-594.

11. Stoner, J.D., D.L. Lorenz, G.J. Wiche, and R.M. Goldstein. 1993. Red River of

the North basin, Minnesota, North Dakota, and South Dakota. American Water

Resources Association 29(4):575-615. 107

12. Wallus, R., and J.P. Buchanan. 1989. Contributions to the reproductive biology

and early life ecology of the Mooneye in the Tennessee and Cumberland River.

The American Midland Naturalist 122(1):204-207.

108

CHAPTER 8

FAMILY ANGUILLIDAE

Introduction

Anguillid eels, or Freshwater Eels, are one of the most unique and widely dispersed groups of fishes in the world, given their catadromous (migrating from freshwater to saltwater to spawn) life history. Although widely dispersed, they are also one of the most understudied groups of fishes. Similar to other eels within the order

Anguilliformes, Anguillids display an elongate, snake-like body shape with the dorsal, caudal, and anal fins conjoined into a single continuous and soft-rayed fin that extends around the posterior end of the body. Small pectoral fins are also present on the anterior end of the body near the head, however any pelvic fins are absent. The family

Anguillidae comprises one genus, Anguilla, with 16 species occurring worldwide, and only one species occurring in North America, the American Eel, Anguilla rostrata.

Like other freshwater eels around the world, the American Eel exhibits a unique spawning migration pattern that consists of moving from freshwater to saltwater.

Although exact spawning locations still remain somewhat of a mystery, it was discovered in the 1920’s by the Danish biologist, Johannes Schmidt, that the American Eel and the

European Eel, Anguilla anguilla, specifically target the Sargasso Sea near Bermuda and the West Indes as a primary spawning site. After traveling sometimes extremely long distances to reach the Sargasso Sea, the American Eel likely spawns at depths up to 500m

(1,640ft), although the actual spawning act has never been observed. After hatching, the

American Eel undergoes five main morphological life stages: leptocephalus, glass eel, 109 elver, yellow eel, and silver eel. Once spawning commences, American Eel exhibit their semelparous behavior, meaning they die shortly after spawning.

After hatching, the leptocephalus, the larvae stage of American Eel, drift in the upper portion of the current for up to a year before they reach the coastal rivers and streams in North America. By this time, the leptocephalus turn transparent in color, and are now referred to as glass eels. As the glass eels continue to migrate further into inland freshwater habitats, they completely metamorphosize into a darker pigment where they are referred to as elvers. Once elvers reach roughly 127mm (5in) in length, they turn a yellow-green color, and are referred to as yellow eels, which is the longest of the life stages. After the yellow eels, essentially nonmigratory juveniles, reach sexual maturity, they turn a silver color and are now referred to as silver eels. The silver eels then begin their long migrations back to the Sargasso Sea to spawn.

The majority of American Eels in the United States occur along the Atlantic Coast, however over several years, the species is capable of traveling as far inland as the

Dakotas. In the Dakotas, the American Eel is rarely documented and observed as they occur in low densities and infrequently inhabit the Missouri River below Gavins Point

Dam near Yankton, South Dakota.

110

American eel, Anguilla rostrata (Lesueur, 1817)

Etymology and Synonyms: Anguilla = Latin for “eel”; rostrata = “beaked” or “long nosed” in reference to the elongate and slim look of the head and body.

Description: Body elongate, snake-like. Adults dorsally black, dark brown to bronze; laterally brown to tan; ventrally light brown to silver; fins dark brown to bronze with no spots or markings. Head small. Snout moderately pointed. Eyes moderate, placed laterally on head. Mouth large, terminal; upper jaw extends to or just beyond posterior edge of eye. Teeth small, numerous; present on both jaws. Single slit gill opening at base of pectoral fin. Dorsal fin elongate, insertion far behind head, continuous with caudal and anal fins. Caudal fin homocercal, slightly rounded. Anal fin insertion posterior from dorsal fin insertion. Pelvic fins absent. Pectoral fins present, small, insertion behind gill covers. Lateral line complete. Adults with numerous inconspicuous and embedded cycloid scales, not visible to human eye. Larvae, or leptocephalus, are strongly laterally compressed. Glass eels with pigmented eyes and transparent body. Elvers dark in color.

Yellow phase eels yellow-green in color.

Similar Species: May be confused with lamprey species in the Dakotas, however lampreys lack any pectoral fins and jaws, and instead have a disk shaped mouth with multiple small teeth in a circle formation.

Distribution and Habitat: One of the most widely dispersed fish in North America. The species is highly adaptive considering is large geographical range that extends from the southern tip of Greenland through North and Central America, and south to Venezuela.28

Inhabits a wide variety of habitat types including streams, lakes, marshes, open oceans, and estuaries with freshwater, marine or brackish water and mud, sand, or gravel 111 substrate.16,28 Although widespread, some populations are in decline due to factors such as overharvesting, contaminants, barriers to migration, and habitat loss and alteration.7,14

Rarely documented and collected in the Dakotas; Gavins Point Dam creates a barrier for movement up the Missouri River.2 Catadromous species. Burrow into soft benthic substrates to lower risk of predation or to escape light and lowering water levels.28

Although adaptable to a wide variety of temperatures, the reported mean temperature preference for yellow eels is 16.7°C (62.06°F).4

Reproduction: Long pre-reproductive migration to saltwater takes place in late summer and early fall, but may begin earlier in well inland lakes.11 Spawning in the southern warmer region of the Sargasso Sea takes place during winter and early spring in the upper

500m (1,640ft) section of the water column.6,20,22,26 Little information is available on the broadcast spawners, and age at sexual maturity not well defined.11 Age and size of males at first reproduction ranges 4-16 years and 228-398mm (8.97-15.67in) TL.13 Age and size of females at first reproduction ranges 6-43 years and 400-1159mm (15.75-45.63in) TL.13

Fecundity of females highly dependent on size.11 Estimates of fecundity from average counts of aliquots for an individual eel ranged 1.84-19.92 million eggs for eels 452-

1133mm (17.80-44.61in) in length.3 Most migrating female eggs <0.25-1.1mm (0.01-

0.04in) in diameter.28,29,30 Low eel density can influence eels to develop into females, and high eel density can influence eels to develop into males.21,24 Hatching presumed to take place February-April, or when eggs are able to incubate 3-4 days at 20°C (68°F).11,13,20

Semelparous species; death occurs shortly after spawning. American eels go through 5 main life stages: leptocephalus, glass, elver, yellow, and silver.12,28 After hatching, not parental care is provided and the larvae, called leptocephalus, drift with the current in the 112 upper portion of the water column for several months until they reach coastal rivers and streams.6,19 At this point leptocephalus turn transparent in color and are now referred to as

“glass eels”, which continue to migrate inland.6,9 Glass eels then morph into dark pigmented “elver”, which continue to seek out migrate farther up into freshwater habitats with their acute sense of smell.11 Elvers >127mm (5in) in length turn yellow-green in color and enter the “yellow phase”, which is the longest life stage lasting many years.11,28

Silver life stage is reached following sexual maturation when the eels begin to migrate back downriver to spawn in saltwater.11

Age and Growth: Length at hatching unknown.11 Growth rate of leptocephali estimated as 0.243mm (0.01in)/day.32 Growth of elvers is slow, reaching roughly 65-127mm (2.56-

5in) in length after the first year in freshwater.5,11,27 Majority of growth occurs during the yellow-phase.11,28 Growth rates highly variable within year classes, leading to poor predictability of length-at-age.11 Adult, or silver females grow to larger sizes than males.11 Mean and maximum size of females increases with latitude.10,16,17 Along the

North American Atlantic Coast, male and female growth rates are inversely correlated to latitude.18,25 Usually 609.6-914.4(24-36in) in South Dakota waters.8 Capable of reaching

1520mm (5ft) in length.1 Longevity of males is 15 years; longevity of females is 43 years.10,13,17

Food and Feeding: Voracious carnivores. Contrary to popular belief, eels prefer to forage on living organisms, and do not often consume dead or rotting prey. Foraging primarily takes place at night. Eels use three main modes of feeding: suction, shaking, and spinning.6,15 Size of prey items increase with size of eel.31 Juvenile life stages 113 primarily feed on larval mayflies, stoneflies, and caddisflies.23 Adults consume a variety of fish, crayfish, invertebrates and bivalves.

Literature Cited:

1. Angel, N.B., and W.R. Jones. 1974. Aquaculture of the American eel (Anguilla

rostrata). International Extension Service School of Engineering, North Carolina

State University.

2. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

3. Barbin, G.P., and J.D. McCleave. 1997. Fecundity of the American eel Anguilla

rostrata at 45°N in Maine, USA. Journal of Fish Biology 51:840-847.

4. Barila, F.Y., and J.R. Stauffer, Jr. 1980. Temperature behavioral responses of the

American eel, Anguilla rostrata (LeSueur), from Mayrland. Hydrobiologia 74:49-

51.

5. Bigelow, H.B., and W.C. Schroeder. 1953. Fishes of the Gulf of Maine. U.S. Fish

and Wildlife Service Fisheries Bulletin 53.

6. Braham, M.A. 2012. Selection of benthic habitat by yellow-phase American eels

(Anguilla rostrata). M.S. Thesis, West Virginia University, Morgantown.

7. Castonguay, M., P.V. Hudson, C.M. Couillard, M.J. Eckersley, J.D. Dutil, and G.

Verreault. 1994. Why is recruitment of the American eel, Anguilla rostrata,

declining in the St. Lawrence River and Gulf? Canadian Journal of Fisheries and

Aquatic Sciences 51:479-488.

8. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls. 114

9. Dutil, J.D., P. Dumont, D.K. Cairns, P.S. Galbraith, G. Verreault, M. Castonguay,

and S. Proulx. 2009. Anguilla rostrata glass eel migration and recruitment in the

estuary and Gulf of St. Lawrence. Journal of Fish Biology 74:1970-1984.

10. Facey, D.E., and G.W. LaBar. 1981. Biology of American eels in Lake

Champlain, Vermont. Transactions of the American Fisheries Society 110:396-

402.

11. Facey, D.E., and M.J. Van Den Avyle. 1987. Species profiles: life histories and

environmental requirements of coastal fishes and invertebrates (north Atlantic):

American eel. U.S. Fish and Wildlife Service Biological Report 82(11.74). U.S.

Army Corps of Engineers, TR EL-82-4.

12. Fahay, M.P. 1978. Biological and fisheries data on American eel, Anguilla

rostrata (leSueue). Technical Series Report No.17. National Marine Fisheries

Service, NOAA, Highlands, New Jersey.

13. Haro, A. 2014. Anguillidae: freshwater eels. Pages 313-331 in M.L. Warren, Jr.,

and B.M. Burr (editors). Freshwater fishes of North America. Johns Hopkins

University Press, Baltimore.

14. Haro, A.J., W. Richkus, K. Whalen, A. Hoar, W.D. Busch, S. Lary, T. Brush, and

D. Dixon. 2000. Population decline of the American eel: implications for research

and management. Fisheries 25:7-16.

15. Helfman, G.S., and J.L. Clark. 1986. Rotational feeding: overcoming gape-limited

foraging in anguillid eels. Copeia 1986:679-685. 115

16. Helfman, G.S., D.E. Facey, L.S. Jr. Hales, and E.L. Jr. Bozeman. 1987.

Reproductive ecology of the American eel. American Fisheries Society

Symposium 1, 42-56.

17. Hurley, D.A. 1972. The American eel (Anguilla rostrata) in eastern Ontario.

Canadian Journal of Fisheries and Aquatic Sciences 29:535-543.

18. Jessop, B.M. 2010. Geographic effects on American eel (Anguilla rostrata) life

history characteristics and strategies. Canadian Journal of Fisheries and Aquatic

Sciences 67:326-346.

19. Kleckner, R.C., and J.D. McCleave. 1982. Entry of migrating American eel

leptocephali into the Gulf Stream system. Helgolander Meeresuntersuchungen

35:329-339.

20. Kleckner, R.C., J.D. McCleave, and G.S. Wippelhauser. 1983. Spawning of

American eel, Anguilla rostrata, relative to thermal fronts in the Sargasso Sea.

Environmental Biology of Fishes 9:289-293.

21. Krueger, W.H., and K. Oliveira. 1999. Evidence for environmental sex

determination in the American eel, Anguilla rostrata. Environmental Biology of

Fishes 55:381-389.

22. McCleave, J.D., R.C. Kleckner, and M. Castonguay. 1987. Reproductive

sympatry of American and European eel and implications for migration and

taxonomy. American Fisheries Society Symposium 1:268-297.

23. Ogden, J.C. 1970. Relative abundance, food habits and age of the American eel,

Anguilla rostrata, in certain New Jersey streams. Transaction of the American

Fisheries Society 99:54-59. 116

24. Oliveira, K. 1999. Life history characteristics and strategies of the American eel,

Anguilla rostrata. Canadian Journal of Fisheries and Aquatic Sciences 56:795-

802.

25. Oliveira, K., and J.D. McCleave. 2002. Sexually different growth histories of the

American eel in four rivers of Maine. Transactions of the American Fisheries

Society 131:203-211.

26. Schmidt, J. 1923. The breeding places of the eel. Philosophical Transactions of

the Royal Society of London Series B 211:179-208.

27. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

28. Tesch, F.W. 1977. The Eel. Biology and management of anguillid eels. Chapman

and Hall, London.

29. Todd, P.R. 1981. Morphometric changes, gonad histology, and fecundity

estimates in migrating New Zealand freshwater eels. New Zealand Journal of

marine and Freshwater Research 15:155-170.

30. Wenner, C.A, and J.A. Musick. 1974. Fecundity and gonad observations of the

American eel, Anguilla rostrata, migrating from Chesapeake Bay, Virginia.

Journal of the Fisheries Research Board of Canada 31:1387-1391.

31. Wenner, C.A., and J.A. Musick. 1975. Food habits and seasonal abundance of the

American eel, Anguilla rostrata, from the Lower Chesapeake Bay. Chesapeake

Science 16:62-66.

32. Wippelhauser, G.S., J.D. McCleave, and R.C. Kleckner. 1985. Anguilla rostrata

leptocephali in the Sargasso Sea during February and March 1981. Dana 4:93-98. 117

CHAPTER 9

FAMILY CLUPEIDAE

Introduction

The Herring family, Clupeidae, consists of roughly 180 species and is primarily made of marine derived species that exhibit anadromous behavior. However, a few species, including the four that occur within the Dakotas, can live their entire lives in inland freshwaters. Due to their similar appearance, Herrings are most often times mistaken with members of the Mooneye family (Hiodontidae), which also occur in the

Dakotas. Herrings can be easily distinguished from members of the Mooneye family by a short or completely absent lateral line, and a scaled keel along the ventral side of the body, which members of the Mooneye family lack. The dorsal fin insertion on Herrings is also distinctly positioned anterior to the insertion of the anal fin, unlike the Mooneyes where the dorsal and anal fin insertions nearly align. Herrings are rather simple in their overall appearance, as they are generally silver and laterally compressed. Similar to members of the trout family, Salmonidae, Herrings have a small, triangular shaped flap called an axillary process near the base of the pelvic fins. Herrings also have transparent partial membranes that cover the anterior and posterior corners of the eye called adipose eyelids.

Herring that inhabit either marine or freshwater habitats mostly exhibit the same schooling behavior within deep open water. The four species that occur in the Dakotas occur in the open areas within the Missouri River and its major tributaries and reservoirs.

Herrings tend to spawn near the surface of the water relatively close to shorelines and 118 generally have extremely high fecundity. Parental care is not provided at any stage of life, the majority of eggs from the family are pelagic, and all Herrings are subject to high mortality rates during their early stages of life. Diets of Clupeids largely consist of plankton, however some species also rely on smaller fish. In the Dakotas, the Gizzard

Shad, Dorosoma cepedianum, Threadfin Shad, Dorosoma petenense, and the Alewife,

Alosa pseudoharengus, primarily rely on algae, plankton and small aquatic insects, whereas the Skipjack Herring, Alsoa chrysochloris, are primarily piscivorous. The relative abundance of Herring within a population is often known to be determined by the species feeding habits and intensity.

Clupeids are one of the world’s most economically important and heavily exploited families of commercial fishes. They also serve as important forage fish in both marine and freshwater habitats. For example, in the Dakotas, Gizzard Shad are an important component to the diets of Walleye, Sander vitreus, and Largemouth Bass,

Micropterus salmoides. However, they can also negatively affect populations of these species when they reach larger sizes. In certain instances when Gizzard Shad become too large to be predated, they become a nuisance and can quickly drive the abundance of zooplankton to low levels, which affect the recruitment of other sport fish that rely on zooplankton during their life stages, such as Bluegill, Lepomis macrochirus.

119

Alewife, Alosa pseudoharengus (Wilson, 1811)

Etymology and Synonyms: Alosa = an old Saxon name for European Shad, Alosa alosa

; pseudoharengus = “false herring”.

Description: Body fusiform, moderately deep, and strongly laterally compressed.

Dorsally gray to olive; laterally silver with a dark, diffuse blue-black spot (similar in size to the pupil) located posteriorly from the upper edge of the gill cover; ventrally silver to white; dorsal, caudal and pectoral fins with gray coloration; anal and pelvic fins transparent to white. Head small. Snout pointed, not protruding beyond upper jaw. Eye large, placed laterally on head. Mouth large, superior, strongly oblique; upper jaw extends to middle of eye; lower jaw protrudes beyond upper jaw. Frenum absent. Barbels absent. Gill rakers fine, 30-44 on first arch. Dorsal fin with 13-14 rays, with posterior ray short and not extending into a long filament. Adipose fin absent. Caudal peduncle short, thick. Caudal fin forked. Anal fin with 17-18 rays and straight distal end. Pelvic fins abdominal with 9 rays; insertions inferior to slightly posterior of dorsal fin insertion.

Lateral line absent with 42-50 cycloid scales in series. Scales present on predorsal midline. Ventral keel with 17-20 scaled scutes anterior to pelvic fin, and 12-15 posterior of pelvic fin.

Similar Species: Resembles the Skipjack Herring. Skipjack Herring have a more elongated and conical head and snout, along with small, but visible teeth on jaws, and a dorsal fin with 17 rays. Gizzard Shad have a more bulbous and fleshy snout that protrudes beyond the upper jaw, a dorsal fin with the last ray extended into a long filament, and have an anal fin with 29 or more rays. Threadfin Shad have fins with a 120 yellow to gold tint, a dorsal fin with the last ray extended into a long filament, and lack scales on the predorsal midline.

Distribution and Habitat: Anadromous species native to the Atlantic Coast drainages from Red Bay, Labrador to South Carolina.6 Has invaded and been introduced to many landlocked waterbodies including the Great Lakes.6,8 Does not occur within North

Dakota. Occurs within the Missouri River below Fort Randall Dam in South Dakota.

Inhabits lentic waters of large rivers, streams, lakes and reservoirs. Larvae and young-of- the-year congregate in large schools in shallow, warmer shoreline areas until early fall, when they begin to migrate towards deeper water.13 Adults exhibit diel migrations, moving to deeper depths during daylight and closer towards the surface at night.14 The critical lower lethal temperature of mature alewives is roughly 3°C (37.4°F).13,15 The critical thermal maximum of the species increases with acclimation temperature for both young-of-the-year and mature individuals, and it is believed that young-of-the year are more tolerant of higher water temperatures than mature individuals.13 The ultimate upper lethal temperature is estimated to be 31-34°C (87.8-93.2°F), when acclimated to 27°C

(80.6°F).7 Tolerant of a wide range of salinities and suspended sediments.14

Reproduction: Spawning takes place during late spring and summer in a variety of habitats including small streams, large rivers, lakes and ponds over a wide range of substrates.1,9,13,14 Sexual maturity may be reached at age-1, however the majority of spawning individuals are within age-2 and age-3.9 It is suggested that Alewives spawn but once.9 Adults migrate into shallow, inshore waters to spawn where they will remain until mid-summer before they migrate back towards deeper waters.9,13,15 Spawning behavior peaks around midnight to the early morning, and consists of two or more 121 individuals swimming rapidly near the surface in a tight circular motion close together with their lateral sides touching.1,2 Females from Lake Michigan produce 11,000-22,000 eggs, however anadromous females from Chesapeake Bay are capable of producing

60,000-100,000 eggs.4,9,14 Eggs initially are demersal and adhesive, and become pelagic and nonadhesive after water hardening.14 Eggs pale yellow in color and roughly 1.0 mm

(0.04 in) in diameter.10 Hatching occurs in water 6.88-29.39°C (44.4-84.9°F), with an optimum hatching temperature of 17.77°C (64°F).3 Eggs hatch at 15 days in water 7.22°C

(45°F), 3.7 days at 21.11°C (70°F), and 2.1 days at 28.89°C (84°F).3

Age and Growth: Larvae roughly 3.8-4.3 mm (0.15-0.17 in) TL at hatching.9,10 Majority of growth achieved within the first year of life.9 Mean lengths-at-age of individuals from

Lake Michigan are reported as: age-0, 96.3 mm (3.79 in) TL; age-1, 138.3 mm (5.44 in)

TL; age-2, 158 mm (6.22 in) TL; age-3, 172.8 mm (6.80 in) TL; age-4, 192.5 mm (7.58 in) TL.9 Capable of reaching 350 mm (13.78 in).6 Longevity 10 years.12,14

Food and Feeding: Diet similar to that of the Gizzard Shad. Juveniles primarily consume large amounts of microcrustacea such as copepods and cladocerans.11 Adults continue to consume large amounts of microcrustacea, but are also known to feed on chironomid larvae, filamentous algae, gastropods and fish eggs of their own kind.1,2,5 Feeding activity is known to be reduced during the spawning migration season.14

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Edsall, T.A. 1964. Feeding by three species of fishes in eggs of spawning

alewives. Copeia 1964:226-227. 122

3. Edsall, T.A. 1970. The effect of temperature on the rate of development and

survival of alewife eggs and larvae. Transactions of the American Fisheries

Society 99:376-380.

4. Foerster, J.W., and S.L. Goodbred. 1978. Evidence for a resident alewife

population in the northern Chesapeake Bay. Estuarine and Coastal Marine

Science 7:437-444.

5. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

6. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

7. McCauley, R.W., and F.P. Binkowski. 1982. Thermal tolerance of the alewife.

Transactions of the American Fisheries Society 111:389-391.

8. Miller, R.R. 1957. Origin and dispersal of the alewife, Alosa pseudoharengus, and

the gizzard shad, Dorosoma cepedianum, in the Great Lakes. Transactions of the

American Fisheries Society 86:97-111.

9. Norden, C.R. 1967a. Age, growth and fecundity of the alewife, Alosa

pseudoharengus (Wilson), in Lake Michigan. Transactions of the American

Fisheries Society 96:387-393.

10. Norden, C.R. 1967b. Development and identification of the larval alewife, Alosa

pseudoharengus (Wilson), in Lake Michigan. Proceedings of the Conference

Great Lakes Research 10:70-78. 123

11. Norden, C.R. 1968. Morphology and food habits of the larval alewife, Alosa

pseudoharengus (Wilson), in Lake Michigan. Proceedings of the Conference

Great Lakes Research 11:103-110.

12. O’Neill, J.T. 1980. Aspects of the life histories of anadromous alewife and the

blueback herring, Marqaree River and Lake Aimsle, Nova Scotia, 1978-1979.

M.S. Thesis, Acadia University, Wolfville, Nova Scotia, Canada. 306pp.

13. Otto, R.G., M.A. Kitchell, and J.O. Rice. 1976. Lethal and preferred temperatures

of the alewife (Alosa pseudoharengus) in Lake Michigan. Transactions of the

American Fisheries Society 105:96-106.

14. Pardue, G.B. 1983. Habitat suitability index models: alewife and blueback

herring. U.S. Department of the Interior Fish and Wildlife Service. FWS/OBS-

82/10.58. 22pp.

15. Stanley, J.G., and P.J. Colby. 1971. Effects of temperature on electrolyte balance

and osmoregulation in the alewife (Alosa pseudoharengus) in fresh and sea water.

Transactions of the American Fisheries Society 100:624-638.

124

Gizzard Shad, Dorosoma cepedianum (Lesueur 1818)

Etymology and Synonyms: Dorosoma; doris = lance, from the Latin word ‘lanceolate’, referring to the shape of the head; soma = ‘body’, referring to the body shape; cepedianum = patronymic for the French ichthyologist Bernard Germain Etienns

LaCepede. The common name, Gizzard Shad, stems from the gizzard-like stomach which aids in the breakdown of food.

Description: Body deep and laterally compressed on each side. Dorsal side silvery blue in color fading to silver laterally with a dark purple to black spot larger than the pupil behind the upper portion of gill cover. This dark spot may be lacking in older individuals.

Head small and scaleless. Snout bulbous, fleshy; protrudes beyond the upper jaw. Mouth small and subterminal containing a small notch in the middle of the premaxilla; upper jaw extends past lower jaw. Frenum absent. Barbels absent. Teeth absent in adults, however teeth are present in larval fish. Eyes large with adipose eyelids present. Gill rakers long and fine, and increase in number with age. Dorsal fin with 10-15 rays; last ray of dorsal fin elongates into a filament extending halfway back to the base of the caudal fin; insertion in the middle of the body, posterior to the insertion of the pelvic fins. Caudal fin forked. Anal fin elongate with 29-37 rays and straight distal end. Pelvic fins abdominal with 7-10 rays. Pectoral fins with 12-17 rays. Lateral line absent with 52-70 cycloid scales in series. Ventral keel is serrated with 17-20 hardened scaled scutes anterior of the pelvic fin, and 10-14 posterior of the pelvic fin. Gizzard Shad have no external sexual dimorphisms.

Similar Species: Closely resembles the Threadfin Shad. Threadfin Shad have a more pointed snout that does not protrude past the upper jaw, have an anal fin with 17-25 rays, 125 and have a lesser number of ventral keel sclaes. Mooneye and Goldeye both possess a lateral line, a dorsal fin base that is generally over the anal fin, and a toothed tongue unlike the Gizzard Shad. Skipjack Herring and Alewife have a lower jaw extending beyond the upper jaw when the mouth is closed, and have a dorsal fin that lacks a long filament. Juvenile Gizzard Shad may be confused with Silver Carp and Bighead Carp, although both species have a complete lateral line.

Distribution and Habitat: Gizzard Shad are native to North America from New Mexico to New York, and from southern Ontario and Quebec to northern Mexico.10,19 In the

Dakotas, the species is primarily found throughout the Missouri and James river basins.

Gizzard Shad are also present in the Bad, Cheyenne, Belle Fourche, and Grand River basins of western South Dakota, the Big Sioux and Vermillion river basins of eastern

South Dakota, and the Cannonball and Heart river basins of western North Dakota.

Gizzard Shad are most numerous in lakes, oxbows, impoundments, and larger streams with low gradients.15 In the Dakotas, Gizzard Shad are often found over mud and fine silt substrates, with adults most frequently occurring in deeper water, and juveniles most often found swimming in schools near the surface of open waters.11 Sudden changes in oxygen and temperature levels can cause mortality, which explains why this species is prone to rapid winter-kill.15

Reproduction: Some females and most male Gizzard Shad mature by age-1, however the majority of females mature by age-2 or 3.1 Spawning occurs in late spring and early summer at temperatures between >10 °C (50 °F) and 21 °C (70 °F).11,15 Spawning takes place during the day in shallow open water bays and inlets often near the surface, and sometimes at night.11,15 Gizzard Shad have also been known to migrate up into smaller 126 streams to spawn.3 Spawning takes place near the surface of the water when mature males and females school together, and swim, roll, and tumble while expelling eggs and milt.14

The fertilized eggs are demersal, and attach to vegetation or other debris.16 Eggs are roughly 0.75 mm (0.03 in) in diameter, and females have a fecundity of roughly 300,000 eggs.1,7,11,16 Since the adults spawn near the surface, no nests are built and no parental care is given.4,5

Age and Growth: Length at hatching is 3.25-5 mm (0.13-0.2 in) TL.18 Early growth is rapid for the species; at age-1 fish reach 101-178 mm (4-7 in) TL, and age-2 fish reach roughly 279 mm (11 in) TL.2,7,11,19 After age-2, female growth is slightly more rapid than that of males.1,11 Average TL is typically larger in northern waters.8,19 In South Dakota,

Gizzard Shad from Angostura Reservoir in the western half of the state attained greater average lengths at each age than individuals from Lake Francis Case and Lake Sharpe.19

Mean lengths-at-age from Angostura Reservoir are reported as: age-3, 381 mm (15 in)

TL; age-6, 447 mm (17.6 in) TL; age-10, 441 mm (17.4 in) TL.19 Capable of reaching

521 mm (20.5 in) TL.17 Gizzard Shad are characterized as short-lived species with high natural mortality rates.9 Longevity 10 years.19

Food and Feeding: Gizzard Shad are an opportunistic omnivorous species. A larval

Gizzard Shad possess teeth during the first summer, and feeds primarily on zooplankton, but are also known to consume phytoplankton, detritus and microcrustaceans.10,11,12

Larvae feed throughout the day and do not feed at night, suggesting that Gizzard Shad larvae are likely visual, particulate feeders.6 Adults consume large amounts of phytoplankton, zooplankton, and detritus. When their intestines become packed, the content is then absorbed by the villi and worked through the gizzard-like stomach where 127 it is digested.3,10 In certain cases, Gizzard Shad are capable of driving zooplankton abundance to low levels, affecting the recruitment of some sport fish.13 One example of sport fishes negatively affected by low abundance of zooplankton are Bluegill and

Largemouth Bass.13 Bluegill rely heavily on zooplankton which in turn affects

Largemouth Bass because Bluegill are the main prey species of Largemouth Bass.13

Literature Cited:

1. Bodola, A. 1965. Life history of the gizzard shad, Dorosoma cepedianum

(Lesueur), in western Lake Erie. Fishery Bulletin 65:391-425.

2. Carlander, K.D. 1969. Handbook of freshwater fishery biology, volume 1. Iowa

State University Press, Ames.

3. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

4. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

5. Cross, F.B., and J.T. Collins. 1995. Fishes in Kansas, second edition, revised.

University of Kansas Natural History Museum, Public Education Series Number

14:1-315.

6. Dettmers, J.M., and R.A. Stein. 1992. Food consumption by larval Gizzard Shad:

zooplankton effects and implications for reservoir communities. Transactions of

the American Fisheries Society 121:494-507.

7. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville. 128

8. Fagan, J.A., and L.C. Fitzpatrick. 1978. Allocation of secondary production to

growth and reproduction by Gizzard Shad Dorosoma cepedianum (Clupeidae) in

Lewisville, Texas. Southwestern Naturalist 23:247-262.

9. Heidinger, R.C. 1983. Life history of gizzard shad and threadfin shad as it relates

to the ecology of small lake fisheries. Pages 1-18 in Proceedings of the Small

Lakes Management Workshop: pros and cons of shad. Iowa Conservation

Commission, Des Moines.

10. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

11. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

12. Megrey, B.A. 1980. Dorosoma cepedianum (Lesueur), gizzard shad. Page 69 in

D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer Jr. editors. Atlas of North American freshwater fishes. North Carolina

State Museum of Natural History, Raleigh.

13. Michaletz P.H. 1998. Population characteristics of Gizzard Shad in Missouri

reservoirs and their relation to reservoir productivity, mean depth, and sport fish

growth. North American Journal of Fisheries Management, 18(1): 114-123.

14. Miller, R.R. 1960. Systematic and biology of the Gizzard Shad (Dorosoma

cepedianum) and related fishes. U.S. Fish and Wildlife Service Fisheries Bulletin

60:371-392. 129

15. Owen, J.B. and G.W. Russell. 1981. Distribution of Fishes in North and South

Dakota Basins affected by the Garrison Diversion Unit. University Press of

University of North Dakota, Grand Forks.

16. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City

17. Trautman, M.B. 1981. The fishes of Ohio. Ohio State University Press. 782pp.

18. Williamson, K.L., and P.C. Nelson. 1985. Habitat suitability index models and

instream flow suitability curves: Gizzard Shad. U.S. Fish and Wildlife Service

Biological Report 82(10.112). 33pp.

19. Wuellner, M.R., Graeb, B.D.S., Ward, M.J., and Willis, D.W. 2009. Review of

Gizzard Shad population dynamics at the northwestern edge if its range.

American Fisheries Society Symposium, 62:37-653.

20. Yako, L.A., J.M. Dettmers, and R.A. Stein. 1996. Feeding preferences of

omnivorous Gizzard Shad as influenced by fish size and zooplankton density.

Transactions of the American Fisheries Society 125:753-759.

130

Skipjack Herring, Alosa chrysochloris (Rafinesque, 1820)

Etymology and Synonyms: Alosa = an old Saxon name for European Shad, Alosa alosa; chrysochloris = Greek for “golden green” referring to the dorsal side coloration.

Description: Body fusiform, strongly laterally compressed. Dorsally olive to dusky dark blue; laterally silver to bronze with dark, round spot absent near posterior edge of the gill cover; ventrally silver to white; dorsal, caudal and anal fins with dusky gray tint; pelvic and pectoral fins clear to lightly pigmented. Head large, conical. Snout pointed, not protruding beyond upper jaw. Eyes large, placed laterally on head. Mouth large, terminal; upper jaw extends to middle of eye; tip of lower jaw protrudes well beyond tip of upper jaw. Frenum absent. Barbels absent. Teeth small, present on both jaws and tongue; present in all ages. Gill rakers 18-30 on first lower arch. Dorsal fin with 17-20 rays with posterior ray short and not extending into a long filament. Adipose fin absent. Caudal peduncle short, thick. Caudal fin forked. Anal fin with 17-21 rays and straight to slightly concave distal end. Pelvic fins abdominal, insertions inferior or slightly posterior to insertion of dorsal fin. Lateral line absent with 51-60 cycloid scales in series. Ventral keel with 17-19 scaled scutes anterior to pelvic fin, and 14-17 posterior of pelvic fin.

Similar Species: Resembles the Alewife. Alewife have a slightly deeper body, a dorsal fin with 13-14 rays, and only 42-50 cycloid scales in lateral line series. Threadfin Shad have a smaller head, a dark black, round spot (similar in size to the pupil) located posteriorly from the upper edge of the gill cover, a dorsal fin with the last ray extended into a long filament, and 40-48 cycloid scales in the lateral line series. Gizzard Shad have a more bulbous and fleshy snout that protrudes beyond the upper jaw, a dark black, round spot (similar in size to the pupil) located posteriorly from the upper edge of the gill cover, 131 an anal fin with 29 or more rays, and a dorsal fin with the last ray extended into a long filament.

Distribution and Habitat: Native to the Mississippi River basin from eastern South

Dakota and central Minnesota in the north, east throughout the Ohio River drainage to western Pennsylvania, and south throughout the Gulf Slope drainages to the Gulf of

Mexico.8 Dams and impoundments have negatively affected the Skipjack Herrings distribution and abundance throughout the upper Mississippi and Missouri River basins, by impeding with the species migration patterns, considering it is a very active and migratory fish.2,5,10 Although the species is still uncommon within the upper Mississippi and Missouri River systems, dredging and other modifications implemented to decrease turbidity within the middle Missouri River have made the species occur more frequently upstream.5,8,9,10 The species is most abundant in the upper Mississippi River below the

Ohio River confluence.10 Not reported in North Dakota. Occurs as far north as Fort

Randall Dam in the Missouri River, as well as the Vermillion River confluence, and the

Whetstone River in the Minnesota watershed.7 Previously reported from Big Stone Lake, but is now likely extripated.1 Often found congregating in large schools within clear to moderately turbid, deep, swift, and open waters of large rivers and occasionally reservoirs.6,8,10,11 Generally avoids and is intolerant of continuous highly turbid waters.10,11

Reproduction: Information on the reproductive habits of the Skipjack Herring is scarce.

Spawning season is prolonged, and likely takes place April through July in the upper

Mississippi River.4 Sexual maturity typically reached at ages 2-3, or when individuals reach roughly 254-305 mm (10.00-12.00 in) TL.6,3 Spawning takes place in the main 132 channel of large rivers over sand and gravel bars.3 Spawning behavior has not been observed although is it presumed that individuals do not spawn in large aggregations.2,3,4

Fecundity dependent on the size of the female, but may range from 76,000-962,000 eggs annually.3 Eggs roughly 0.8-1.1 mm (0.03-0.04 in) in diameter.2,3 Hatching takes place within 34-48 hours at a water temperature of 17.2°C (62.69°F).3

Age and Growth: Young-of-the-year generally reach 75-150 mm (2.95-5.91 in) TL by the end of their first year.10,6 In Ohio, young-of-the-year are reported to reach 25-100 mm

(0.98-3.94 in) in August and 130-200 mm (5.12-7.87 in) in October.11 Capable of reaching 533 mm (20.98 in) TL.11 Longevity 4 years.3

Food and Feeding: Piscivores. Individuals >40 mm (1.57 in) TL primarily forage on zooplankton, and juveniles consume zooplankton, insect larvae such as dipterans, and small fish.3 Adults consume a greater amount of fish as they increase in size, and have been known to consume Gizzard Shad and Threadfin Shad near the surface.6 Feeding behavior also includes large schools of individuals congregating in swift waters where they force large schools of minnows, such as Emerald Shiner, near the water’s surface.11

Once the prey is near the surface, Skipjack Herring will dash into the school of minnows, and may even leap out of the water to capture jumping minnows.11 This jumping or

“skipping” motion is likely what gave the species its common name.11

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 133

3. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

4. Coker, R.E. 1930. Studies of common fishes of the Mississippi River at Keokuk.

Bulletin of the US Bureau of Fisheries 1072:141-225.

5. Cross, F.B., and D.G. Huggins. 1975. Skipjack herring, Alosa chrysochloris, in

the Missouri River. Copeia 1975:382-385.

6. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

7. Hayer, C.A. 2014. Fish assemblage structure, trophic ecology, and potential

effects of invading Asian carps in three Missouri River tributaries, South Dakota.

Ph.D. Dissertation, South Dakota State University, Brookings, South Dakota.

8. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

9. Neebling, T.E., and M.C. Quist. 2008. Observations on the distribution and status

of western sand darter, spotted gar, and skipjack herring in Iowa rivers. The

Journal of the Iowa Academy of Science 115:24-27.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

11. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

134

Threadfin Shad, Dorosoma petenense (Günther, 1867)

Etymology and Synonyms: Dorosoma = Greek for “lanceolate body”, referring to the body shape of juveniles; petenense = named after the type locality, Lake Peten, Yucatan.

Description: Body fusiform, moderately deep, laterally compressed. Dorsally olive to dusky dark blue; laterally silver with a dark black, round spot (similar in size to the pupil) located posteriorly from the upper edge of the gill cover; present in all ages; ventrally silver to white; all fins except the dorsal fin with yellow to gold tint. Head small. Snout pointed, not protruding beyond upper jaw. Eyes large, placed laterally on head. Mouth large, terminal; upper jaw does not extend beyond middle of eye. Frenum absent. Barbels absent. Gill rakers long, fine and numerous. Dorsal fin with 11-15 rays, with last ray extended into a long filament often reaching to posterior end of the anal fin. Adipose fin absent. Caudal peduncle short, thick. Caudal fin forked. Anal fin elongate with 17-25 rays and straight distal end. Pelvic fins abdominal with 7-8 rays; insertions directly inferior to insertion of dorsal fin. Pectoral fins with 12-17 rays. Lateral line absent with

40-48 cycloid scales in series. Scales absent on predorsal midline. Ventral keel with 16-

17 scaled scutes anterior of pelvic fin, and 9-11 posterior of pelvic fin.

Similar Species: Closely resembles the Gizzard Shad. Gizzard Shad have a more bulbous and fleshy snout that protrudes beyond the upper jaw, an anal fin with 29 or more rays, and fins dusky gray in color, lacking any yellow to gold tint. Alewife lack a long filament in the dorsal fin, and have a scaled predorsal midline.

Distribution and Habitat: Native to the lower Mississippi River and Gulf Slope drainages, south to Central America. Have been widely introduced and stocked outside its native range, particularly in large reservoirs, to act as a forage species for piscivorous 135 game fish and to manipulate predator-prey interactions.3 Does not occur within North

Dakota. Occurs in the Missouri River as far north as Lewis and Clark Lake in South

Dakota. Inhabits reservoirs and big rivers with swift current.4 Intolerant of cold-water temperatures, and often experiences mass mortalities during winter in areas where temperatures suddenly drop or are severe for longer periods of time.6,15 In aquaria, lower lethal temperature was determined to be approximately 5°C (41°F), when individuals were exposed to temperatures decreasing at the rate of 1°C every 72 hours.6

Reproduction: Spawning season is prolonged. Sexual maturity usually reached at sizes

<130 mm (5.12 in).6 Spawning behavior consists of schools splashing and swimming quickly back and forth through beds of vegetative matter near the water’s surface and shoreline.5,13 No nest is constructed and no parental care is given. Spawning lasts for 5-10 seconds before individuals immediately return to deeper water.5 Fecundity of females from central Arizona reservoirs ranged 923-8,540 eggs, but are capable of producing up to 12,400 eggs.11,12 The number of mature ova/ gram of a female not known to be dependent on the length, weight, or age of a female.11 Eggs roughly 0.82 mm (0.03 in) in diameter and adhesive to vegetation.5,11 Maximum developmental temperature for the successful incubation of eggs is reported to be in water temperatures below 34.2°C

(93.56°F).8 Maximum survival rates for cohorts of larvae is greatest at roughly 22°C

(71.6°F), and begins to decease at higher temperatures.1

Age and Growth: Little information is available on growth rates of Threadfin Shad within the Dakotas region. Mean lengths-at-age of individuals from a Salt River reservoir in central Arizona are reported as: age-1, 39.0 mm (1.54 in) SL; age-2, 63.2 mm (2.49 in)

SL; age-3, 78.7 mm (3.10 in) SL; age-4, 92.7 mm (3.65 in) SL.10 Majority of growth 136 takes place over the spring and summer months.10 Growth rates of larvae up to 21 days old are known to be related to water temperature and prey availability.1 Capable of reaching 220 mm (8.66 in) TL.2 Longevity 3 years.2,16

Food and Feeding: Opportunistic sight feeders, with no apparent diet differences in individuals of different sizes.9,14 Although they are primarily considered pelagic planktivores given their long and fine gill filaments to filter out phytoplankton and zooplankton, studies have also revealed that Threadfin Shad can shift their diet to feed extensively along the bottom for prey items, such as chironomids, when their primary diet items are lacking.7,9 Feeding behavior consists of quick movements back and forth with their mouths propped open for roughly 2 or 3 seconds at a time.5 Relies heavily on small crustaceans such as zooplankton, but also known to consume greater amounts of vegetation, organic matter, and other organisms of suitable size.5,7,14 Feeding activity is known to decrease when water temperature reaches roughly 10°C (50°F).6

Literature Cited:

1. Betsill, R.K., and M.J. Van Den Avyle. 1997. Effect of temperature and

zooplankton abundance on growth and survival of larval threadfin shad.

Transactions of the American Fisheries Society 126:999-1011.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. DeVries, D.R., R.A. Stein, J.G. Miner, and G.G. Mittelbach. 1991. Stocking

threadfin shad: consequences for young-of-year fishes. Transactions of the

American Fisheries Society 120:368-381. 137

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

5. Gerdes, J.H., and WM.J. McConnell. 1963. Food habits and spawning of the

threadfin shad in a small, desert impoundment. Journal of the Arizona Academy

of Science 2:113-116.

6. Griffith, J.S. 1978. Effects of low temperature on the survival and behavior of

threadfin shad, Dorosoma petenense. Transactions of the American Fisheries

Society 107: 63-70.

7. Haskell, WM.L. 1959. Diet of the Mississippi threadfin shad, Dorosoma

petenense atchafalayae, in Arizona. Copeia 1959:298-302.

8. Hubbs, C., and C. Bryan. 1974. Maximum incubation temperature of the threadfin

shad, Dorosoma petenense. Transactions of the American Fisheries Society

103:369-371.

9. Ingram, W., and C.D. Ziebell. 1983. Diet shifts to benthic feeding by threadfin

shad. Transactions of the American Fisheries Society 112:554-556.

10. Johnson, J.E. 1970. Age, growth, and population dynamics of Threadfin Shad,

Dorosoma petenense (Günther), in central Arizona reservoirs. Transactions of the

American Fisheries Society 99:739-753.

11. Johnson, J.E. 1971. Maturity and fecundity of threadfin shad, Dorosoma

petenense (Günther), in central Arizona reservoirs. Transactions of the American

Fisheries Society 100:74-85. 138

12. Kilambi, R.V., and R.E. Baglin, Jr. 1969. Fecundity of the threadfin shad,

Dorosoma petenense, in Beaver and Bull Shoals reservoirs. Transactions of the

American Fisheries Society 98:320-322.

13. Lambou, V.W. 1965. Observations and size distribution and spawning behavior of

threadfin shad. Transactions of the American Fisheries Society 94:385-386.

14. Miller, R.V. 1967. Food of the threadfin shad, Dorosoma petenense, in Lake

Chicot, Arkansas. Transactions of the American Fisheries Society 96:243-246.

15. Parsons, J.W., and J.B. Kimsey. 1954. A report on the Mississippi threadfin shad.

The Progressive Fish Culturist 16: 179-181.

16. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

139

CHAPTER 10

FAMILY CYPRINIDAE

Introduction

Cyprinidae is a family of freshwater fishes, commonly called cyprinids, and includes the carps and minnows. Cyprinids are the largest and most diverse family of fish and include well over 2,000 species native to North America, Europe, Asia and Africa.

Cyprinids are well represented in South Dakota, as they make up roughly one third of the fishes that occur throughout both states. Often commonly referred to as the minnow family, many have come to believe that the group encompasses any smaller species of fish, however this grouping based on body size is not accurate. Some smaller species of fish belong to other families, and some larger species such as the Common Carp,

Cyprinus carpio, belong to the cyprinid family. The largest species of cyprinid in the world is the Giant Barb, Catlocarpio siamensis, which is native to Indochina and is capable of reaching 3m (9.8ft) in length, and the largest North American cyprinid is the

Colorado Pikeminnow, Ptychocheilus lucius, which can reach up to 1.8m (6ft) in length.

Cyprinids that are native to the Dakotas are distinguished by having a single dorsal fin with 8-9 rays and no spines, whereas the introduced species of cyprinids like carp, have a spinous dorsal ray, except for the Grass Carp, Ctenopharyngodon idella, which can still be separated from the native cyprinids in the Dakotas by their large, elongate and cylindrical bodies. The spinous dorsal ray on the larger species of cyprinids in the Dakotas helps separate them from the sucker species within both states, from the family Catostomidae. Cyprinids also have internal distinguishing features such as a 140

Weberian apparatus and toothless jaws. The Weberian apparatus is a modified structure of minute bones within the first few vertebrae that connect the swim bladder to the inner ear, to help detect vibrations in the water and help alert the fish from things like predators or structures. Instead of having teeth aligning the jaws, cyprinids have modified gill arches, with the last gill arch having pharyngeal teeth that are arranged in rows and help grind forage material efficiently.

The habitats, reproductive behavior, and diets of cyprinids, present and not present in the Dakotas, are all very diverse among the family. The Northern Pearl Dace,

Margariscus nachtriebi, prefers to inhabit small, coolwater glacial lakes and headwater stream pools of first and second order streams, whereas Sturgeon Chub, Macrhybopsis gelida, prefer to inhabit medium to large turbid rivers, and Golden Shiner, Notemigonus crysoleucas are abundant in lakes and impoundments. Central Stoneoller, Campostoma anomallum, are ubiquitous grazers sweeping their head in a side to side motion along the bottom substrate to scrape algae from rocks using the cartilaginous ridge on the inside of the lower lip. Silver Carp, Hypophthalmichthys molitrix, differ in that they are planktivorous filter feeders, and strain larger amounts of small zooplankton and phytoplankton within the pelagic zone. Fathead Minnow, Pimephales promelas, construct cavity-like nests under rocks, logs or other structures, whereas Flathead Chub, Platygobio gracilis, are broadcast pelagic spawners.

141

Central Stoneroller, Campostoma anomalum (Rafinesque, 1820)

Etymology and Synonyms: Campostoma; campo = curved, stoma = mouth; anomalum

= extraordinary.

Description: Body fusiform, elongate, robust, nearly oval to round anteriorly. Dorsally dark olive to gray; laterally olive to silver with dark mottling, sometimes with faint lateral stripe; ventrally silver to white; fins typically clear with dark bands on dorsal and anal fins on adults; dark dusky spot on caudal fin. Head short, blunt. Snout rounded, slightly protrudes past mouth. Eye tan to amber in color. Mouth subterminal, crescent shaped; maxillary short. Frenum absent. Barbles absent. Lips fleshy; lower lip with inner cartilaginous ridge. Pharyngeal teeth 0,4-4,0; slightly hooked. Gill rakers 21-33. Dorsal fin with 8 rays, no spines. Adipose fin absent. Caudal peduncle moderately thick. Caudal fin slightly forked. Anal fin with 7 rays; insertion posterior to dorsal fin. Pelvic fin abdominal, usually 8 rays; insertion slightly anterior to dorsal fin. Pectoral fin with 15 rays. Lateral line complete; 47-60 cycloid scales; 38-50 circumferential scales. Intestine extremely long and spiraled around air bladder. Peritoneum black. Tubercles on spawning males large, slightly hooked, and white; present on head, body, and median fins. Spawning males also with minor “hump” behind head, and black and orange bands through the central part of dorsal, pelvic, and anal fins. Spawning females may develop a dusky bar within the dorsal fin but remain olive in color. Juveniles with dark, horizontal stripe on sides.

Similar Species: Closely resembles Largescale Stoneroller. Largescale Stoneroller with larger scales, 41-48 in lateral series, and 29-38 circumferencial scales. Juveniles and smaller adults resemble Western Blacknose Dace, which lack a cartilaginous ridge on the 142 lower lip, and also have a frenum present. Common Shiner have elongated, diamond- shaped scales with a height 3.5 times their width, and have fins lacking dark bands.

Smaller White Sucker have thick, protruding lips and a greater number of lateral line scales (53-85). Can be distinguished from other cyprinids by the long, spiraled intestine around the air bladder.

Distribution and Habitat: Native throughout much of the Missouri, Mississippi, and

Ohio River basins. Found scarcely within tributaries of the Red River in North Dakota.

Commonly found in southeastern South Dakota river drainages. Inhabits headwater streams, as well as small to medium sized rivers and streams (sometimes intermittent) with swift, clear water. Rarely found in lakes. Often near riffles, within pools, or along stream banks with overhanging vegetation. Prefers gravel or bedrock substrate. Able to tolerate warmer water temperatures, short-term intermittency, and moderate amounts of siltation and turbidity.8

Reproduction: Spawning migrations to suitable spawning grounds vary with habitat but are known to be upstream movements into small tributaries.10 Spawning grounds consist of shallow areas above riffles with swift current, over fine gravel, or near deep pools with overhanging vegetation. Spawning activity takes place from mid-April to May, and begins when water temperatures reach 18 °C (65 °F).11 Males known to prepare and be territorial of one or more nests by clearing an area surrounded by larger pieces of pebble and gravel which they move with their mouths or snout.10 Nest size known to vary with habitat.10 One or more males escort the female to the nest, where her eggs are fertilized and deposited. No parental care is given, and both males and females retreat back downstream.3 Fecundity 1170-1500 eggs per female.9 Eggs bright yellow, roughly 2.0 143 mm (0.08 in) in diameter, and adhesive to gravel within the nest. Sexual maturity reached at age 1-2.12 Hatching occurs within 2-3 days.

Age and Growth: Newly hatched larvae roughly 5.7 mm (0.22 in) TL.1 Growth rapid within the first 2 years. Males attain greater lengths than females. Length-at-age in

Illinois were calculated as: age-1, 51 mm (2.0 in) TL; age-2, 79 mm (3.1 in) TL; age-3,

99 mm (3.9 in) TL.6 Average 100-177 mm (4-7 in) TL. Capable of reaching 203-254 mm

(8-10 in) TL.1 Larger lengths are reached in the southern portion of their range.

Longevity 3-6 years.9

Food and Feeding: Omnivore. Young feed on rotifers and small zooplankton. Adults primarily consume large amounts of filamentous algae, diatoms and detritus, but also forage on macroinvertebrates, small mollusks and larval fish.2,7,11 Ubiquitous grazers; feed primarily during the day. Feeding behavior noted as moving head from side to side, and scraping attached algae from substrate using the cartilaginous ridge on the inside of the lower lip. Capable of consuming up to 27% of their body weight in benthic algae per day, significantly decreasing algal biomass.3,4,13 Due to this large amount, the species is capable of affecting functional and structural stream ecosystem properties both directly and indirectly.5,8

Literature Cited:

1. Eddy, S., and J.C. Underhill. 1974. Northern fishes with special reference to the

upper Mississippi Valley. Third edition. University of Minnesota Press,

Minneapolis. 414p.

2. Evans-White, M.A., W.K. Dodds, and M.R. Whiles. 2003. Ecosystem

significance of crayfishes and stonerollers in a prairie stream: functional 144

differences between co-occurring omnivores. Journal of the North American

Benthological Society 22:423-441.

3. Fowler, J.F., and C.A. Taber. 1985. Food habits and feeding periodicity in two

sympatric central stonerollers (Cyprinidae). American Midland Naturalist

113:217-223.

4. Gelwick, F.P., and W.J. Matthews. 1992. Effects of an algivorous minnow on

temperate stream ecosystem properties. Ecology 73:1630-1645.

5. Gido, K.B., K.N. Bertrand, J.N. Murdock, W.K. Dodds, and M.R. Whiles. 2010.

Disturbance-mediated effects of fishes on stream ecosystem processes: concepts

and results from highly variable prairie streams. Pages 593-617 in K.B. Gido and

D.A. Jackson (editors). Community ecology of stream fishes: concepts,

approaches, and techniques. American Fisheries Society, Bethesda, Maryland.

6. Gunning, G.E., and W.M. Lewis. 1956. Age and growth of two important bait

species in a cold-water stream in southern Illinois. American Midland naturalist

55:118-120.

7. Hargrave, C.W. 2006. A test of three alternative pathways for consumer

regulation of primary productivity. Oecologia 149:123-152.

8. Matthews, W.J., A.J. Stewart, and M.E. Power. 1987. Grazing fishes as

components of North American stream ecosystems: effects of Campostoma

anomalum. Pages 128-135 in W.J. Matthews and D.C. Heins (editors).

Community and evolutionary ecology of North American stream fishes.

University of Oklahoma Press, Norman, Oklahoma. 145

9. McKee, P.M., and B.J. Parker. 1982. The distribution, biology, and status of the

fishes Campostoma anomalum, Clinostomus elongates, Notropis photogenis

(Cyprinidae), and Fundulus notatus (Cyprinodontidae) in Canada. Canadian

Journal of Zoology 60:1347-1358.

10. Miller, R.J. 1962. Reproductive behavior of the stoneroller minnow, Campostoma

anomallum pullum. Copeia 1962(2):407-417.

11. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

12. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

146

Largescale Stoneroller, Campostoma oligolepis (Hubbs and Greene, 1935)

Etymology and Synonyms: Campostoma = Latin for “curved mouth”, referring to the shape of their jaws which allow for their feeding strategy; oligolepis = Latin for “fewer scales”.

Description: Body fusiform, elongate, moderately robust, oval in cross section. Dorsally gray, brown to dark olive; laterally olive to silver with dark mottling, sometimes with faint lateral stripe; ventrally silver to white; fins typically clear, adults with dark band on dorsal fin; dark dusky spot present on caudal fin. Head short, blunt. Snout rounded, slightly protrudes past upper lip. Eye tan to amber in color. Mouth subterminal, crescent shaped; maxillary short. Frenum absent. Barbles absent. Lips fleshy; lower lip with inner cartilaginous ridge. Pharyngeal teeth 0,4-4,0; slightly hooked. Gill rakers 20-28. Dorsal fin with 8 rays, no spines. Adipose fin absent. Caudal peduncle moderately thick. Caudal fin slightly forked. Anal fin with 7 rays. Pelvic fin abdominal, usually 8 rays; insertion slightly anterior to dorsal fin insertion. Pectoral fin with 13-17 rays, no spines. Lateral line complete; 41-48 cycloid scales; 29-38 circumferential scales. Intestine long, spiraled around swim bladder. Peritoneum black. Tubercles on spawning males large, slightly hooked, and light colored; present on head, body, and median fins; inner-nasal tubercles absent. Spawning males also with small “hump” behind head, red-orange coloring on the ventral scales and pelvic and anal fins, as well as a light pink spot on each side of the snout. Juveniles similar to adults, with dark lateral stripe.

Similar Species: Closely related to and resembles the Central Stoneroller. Central

Stoneroller have a more robust body with smaller scales, 47-60 in lateral series, and 38-

50 circumferencial scales. Anal fin of adult Central Stoneroller with dark band present. 147

Tuburculate Central Stoneroller males with inner-nasal tubercles present. Juvenile

Largescale Stoneroller and smaller adults resemble Blacknose Dace, which lack a cartilaginous ridge on the lower lip, and have a frenum present. Common Shiner have elongated, diamond-shaped scales with a height 3.5 times their width, and have fins lacking dark bands. Smaller White Sucker have thick, protruding lips and a greater number of lateral line scales (53-85). Can be distinguished from other cyprinids by the long, spiraled intestine around the air bladder.

Distribution and Habitat: Native throughout the Upper and Lower Mississippi River basins from northern Wisconsin, into eastern Minnesota and Iowa, and south to the

Mobile basin and part of the Ozark region. Single population known from the Forest

River in northeastern North Dakota. Has not been recorded in South Dakota. Habitat specialist.3,4,8 Adults inhabit headwater streams, as well as small to medium sized rivers and streams with swift current. Often near riffles and swift runs over sand, cobble or gravel substrate. Occasionally found in pools. Known to prefer larger sized riffles and a faster current than the Central Stoneroller.2,4,6 Juveniles favor areas with slower current and sand and silt substrates.2 Adults with a high pollution tolerance, but a low tolerance for silt.2

Reproduction: Due to the Largescale Stoneroller being understudied, information regarding reproduction is often derived from other species within the genus Campostoma.

Spawning known to take place in the spring, with a peak reproductive season from March to April, but may extend into May.10 Peak spawning temperature has been recorded as

12°C, however spawning activity is known to be influenced by environmental variations with season and year.9,10 Fractional spawners.10 Males known to create and defend one to 148 several nests as large as 30cm (12in) by moving pieces of pebble and gravel with their mouths or snout.10 Nest are often near riffles with nearby pools, in shallow waters. One or more males escort a single female to a nest where the eggs are fertilized and deposited.

Once spawning is complete, no parental care is given. Fecundity roughly 1,510 eggs from an age-2 female, 111mm (4.4in) and 14.74g (0.03lb).1 Eggs yellow-amber in color, adhesive, and roughly 1.4mm (0.05in) in diameter.1 Sexual maturity reached at age 1-2.

Age and Growth: Average length 102mm (4 inches).3 Average length at age from the

Eau Claire River in Wisconsin were calculated as: age-1, 55.6mm (2.2in) TL; age-2,

90.3mm (3.5in) TL; age-3, 126.3mm (5.0in) TL.1 Males often reach greater maximum lengths than females.2 Growth rapid within the first year. Capable of reaching 175mm

(6.9in) TL.5 Average longevity 3-5 years.

Food and Feeding: Diet and feeding similar to the Central Stoneroller. Omnivore. Feeds continuously during the day.4 Adults primarily ubiquitous grazers on filamentous algae.

Feeding behavior noted as moving head from side to side and scraping attached algae from substrate using the cartilaginous ridge on the inside of the lower lip. Their long intestine allows the species to consume up to 27% of its body weight in benthic algae per day, significantly decreasing algal biomass.4,7,10 Like the Central Stoneroller, this species is suspected to be capable of creating changes within the ecosystem due to the large amount of algal biomass consumed.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Burr, B.M., and P.W. Smith. 1976. Status of the Largescale Stoneroller,

Campostoma oligolepis. Copeia 1976 (3):521-531. 149

3. Burr, B.M., R.C. Cashner, and W.L. Pflieger. 1979. Campostoma oligolepis and

Notropis ozarcanus (Pisces: Cyprinidae), two additions to the known fish fauna of

the Illinois River, Arkansas and Oklahoma. The Southwestern Naturalist 24:381-

383.

4. Fowler, J.F., and C.A. Taber. 1985. Food habits and feeding periodicity in two

sympatric stonerollers (Cyprinidae). American Midland Naturalist 113(2) 217-

224.

5. Metee, M.F., P.E. O’Neil, and J.M. Pierson. 1996. Fishes of Alabama and the

Mobile Basin. Birmingham, Alabama: Oxmoor House.

6. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

7. Power, M.E., W.J. Matthews, and A.J. Stewart. 1985. Grazing minows,

piscivorous bass, and stream algae: dynamics of a strong interaction. Ecology

66:1448-1456.

8. Rakocinski, C.R. 1977. Evolutionary interactions of two sympatric related species

of minnows in a creek in Northwestern Illinois: hybridization and isolating

mechanisms between Campostoma oligolepis and Campostoma anomalum

pullum. Master’s Thesis, Northern Illinois University, DeKalb, Illinois.

9. South, E.J., and W.E. Ensign. 2013. Life history of Campostoma oligolepis

(Largescale Stoneroller) in urban and rural streams. Southeastern Naturalist

12(4):781-789. 150

10. Timms, D.M. 2017. Reproductive timing of the Largescale Stoneroller,

Campostoma oligolepis, in the Flint River, Alabama. Master’s Thesis, University

of Alabama-Huntsville, Huntsville, Alabama.

151

Goldfish, Carassius auratus (Linnaeus, 1758)

Etymology and Synonyms: Carassius = Latinization of the vernacular name Karass or

Karausche, for the European Crucian carp; auratus = Latin for “gilded”, referring to the golden color.

Description: Body deep, slightly laterally compressed with arched back. Coloration greatly variable; ‘wild’ individuals generally olive or bronze, but may also be solid orange to tangerine, or mottled with white, brown or black blotches. Head short, small, triangular. Snout bluntly pointed. Eye moderately large, placed laterally on head. Mouth moderately small, terminal and oblique. Frenum absent. Barbels absent. Lips thin.

Pharyngeal tooth pattern 0,4-4,0. Gill rakers long, 37-43. Dorsal fin elongate with 1 spinous, serrated soft ray anteriorly, followed by 15-19 rays. Adipose fin absent. Caudal peduncle short, thick. Caudal fin moderately forked. Anal fin with 1 spinous, serrated soft ray anteriorly, followed by 5-6 rays. Pelvic fins abdominal with 8-9 rays; insertions inferior or slightly posterior to insertion of dorsal fin. Pectoral fins with 15-16 rays.

Lateral line complete with 26-31 large cycloid scales in series. Spawning males develop small tubercles on the head, dorsal side of body and on the pectoral fins.

Similar species: Resembles the Common Carp. Common Carp have two pairs of barbels present (1 short barbell on each side of the snout, and 1 longer barbell on each corner of the mouth), 32-41 scales in the lateral line series, and a more conical and elongate head and snout. Bigmouth Buffalo have a large, rounded head, small eyes positioned anteriorly on the head, and pelvic fins with 10-11 rays. Smallmouth Buffalo have a small, conical head creating a steep, or “humped” profile, and a small subterminal mouth with thick lips. 152

Distribution and Habitat: Native to Asia and Japan, but have been widely introduced over the world primarily as an aquarium fish.2 Well established and abundant in some parts of North America, however occurrences are sporadic and are likely due to release or escapement.9 Not reported in North Dakota. In South Dakota, the distribution is limited to

Capitol Lake in Pierre, and the Fall River in Hot Springs, although they have been previously reported within the Cheyenne River drainage and Cody Lake in Bennett county.2 In South Dakota, the species distribution is likely limited due to cold water temperatures since it is considered a warm-water species.2 Mean critical thermal minimum and maximum are known to be significantly related to acclimation temperature.4 When acclimated to 15 °C (59 °F), mean critical thermal minimum is reported at 1.3 °C (34.34 °F), and mean critical thermal maximum is 34.5 °C (94.1 °F).4

When acclimated to 25 °C (77 °F), mean critical thermal minimum is reported at 5.2 °C

(41.36 °F), and mean critical thermal maximum is 39.6 °C (103.28 °F).4 Inhabits heavily vegetated waters with little to no current such as pools in quiet streams, ponds, shallow lakes, and occasionally impoundments.8 Adults are often found near the bottom substrate, but are also known to school near the surface.1 Highly tolerant of a wide range of water temperatures, increased salinity, and low dissolved oxygen levels.5,6,9 Less tolerant of moderate or high gradients, and increased turbidity.13

Reproduction: Spawning season is prolonged, occurring from spring to late summer, and begins when water temperatures reach roughly 15.6 °C (60.08 °F).1 Spawning generally takes place at dawn.5 Males capable of reaching sexual maturity at age-1, and females at age-1 in warmer climates and age-2 in colder climates.5 Multiple males are known to chase a single female into areas with dense vegetation, where intense splashing 153 behavior occurs during which eggs and milt are released.5 No nest is constructed and no parental care is given. Fractional spawners, with females capable of releasing 2,000-

4,000 eggs in a single clutch, and producing 160,000-380,000 eggs total.1 Eggs pale amber in color, adhesive to vegetation, and roughly 1.6 mm (0.06 in) in diameter.1,5,7

Hatching occurs within 46-54 hours at 29 °C (84.2 °F), 5 days at 20 °C (68 °F), and 8-10 days at 15 °C (59 °F).1,10

Age and Growth: Growth of young-of-the-year highly variable.13 Growth dependent on the length of the spawning season, different environmental conditions, and is density dependant.11 Water temperature and salinity are both known to significantly affect growth.6 Mean length-at-age of ‘wild’ individuals from Wisconsin are reported as: age-0,

84.4 mm (3.32 in) TL; age-1, 120.2 mm (4.73 in) TL; age-2, 151.3 mm (5.96 in) TL; and age-3, 188.0 mm (7.40 in) TL.1 Capable of reaching 457 mm (18.00 in) TL.1 Longevity

20 years in captivity, and 30 years in ponds not subjected to severe weather.7

Food and Feeding: Omnivore. Juveniles mainly feed on zooplankton and insect larvae.9

Diets of ‘wild’ Goldfish known to be similar to carp, and primarily consist of diatoms, phytoplankton, small crustaceans, aquatic insects, green algae and aquatic vegetation.3,5,9,12 Spawning adults often consume their own eggs and newly hatched larvae.5

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Blackwell, B.G. 2007. Warm-water fish species. Pages 213-238 in C. Berry, K.

Higgins, D. Willis, and S. Chipps, editors. History of fisheries and fishing in

South Dakota. South Dakota Department of Game, Fish and Parks, Pierre. 154

3. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

4. Ford, T., and T.L. Beitinger. 2005. Temperature tolerance in the goldfish,

Carassius auratus. Journal of Thermal Biology 30:147-152.

5. Horváth, L., G. Tamás, and C. Seagrave. 1992. Carp and pond fish culture:

Including Chinese herbivorous species, pike, tench, zander, wels catfish and

goldfish. Halsted Press, New York.

6. Imanpoor, M.R., E. Najafi, and M. Kabir. 2012. Effects of different salinity and

temperatures on the growth, survival, haematocrit and blood biochemistry of

goldfish (Carassius auratus). Aquaculture Research 43:332-338.

7. Innes, W.T. 1936. The complete aquarium book: The care and breeding of

goldfish and tropical fishes. Halcyon House, New York.

8. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

9. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

10. Mansueti, A.J., and J.D. Hardy. 1967. Development of fishes of the Chesapeake

Bay region. University of Maryland, Baltimore, Natural Resources Institution,

Part I. 202pp.

11. Moyle, P.B. 2002. Inland fishes of California, revised and expanded. University

of California Press, Berkley. 155

12. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

13. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

156

Northern Redbelly Dace, Chrosomus eos (Cope, 1862)

Etymology and Synonyms: Chrosomus = “coloration”; eos = “sunrise”, likely referring to the yellow and orange coloration on males.

Description: Northern Redbelly Dace have a fusiform, elongate and cylindrical body.

Dorsally dark olive to dusky brown; lateral sides with multiple stripes of color beginning with a thin, narrow dark lateral stripe nearest to dorsal side (beginning at posterior edge of operculum and extending to base of caudal fin), then a thicker light olive to silvery-tan lateral stripe below, followed by a thicker dark black lateral stripe (beginning at tip of snout and extending to the base of caudal fin creating a dark caudal spot). Males display a dusky red to yellow-orange stripe below the thicker dark lateral stripe; ventrally silvery white; fins generally clear but may display light peach to cream pigmentation. Head is moderate. Snout blunt. Eye moderately large placed laterally on head. Mouth small, terminal, strongly oblique; lower jaw often protruding past upper jaw; jaws do not extend halfway back towards eye. Lips fleshy. Barbels absent. Pharyngeal teeth slender; pattern

0,5-5,0. Gill rakers short, 10. Dorsal fin has 7-8 rays. Adipose fin absent. Caudal peduncle elongate, thick. Caudal fin moderately forked. Anal fin has 8 rays. Pelvic fins abdominal having 8 rays; insertion distinctly anterior to insertion of dorsal fin. Pectoral fins short, rounded having 13-15 rays. Lateral line incomplete with 70-87 extremely small cycloid scales in series, appearing scaleless with the naked eye. Spawning males develop bright red-orange coloration below the thicker dark lateral stripe, and small tubercles on breast, pectoral fins and caudal peduncle. Juveniles with more dull coloration. 157

Similar Species: Closely resembles Finescale Dace, Southern Redbelly Dace and

Northern Pearl Dace. Finescale Dace display a single distinct dark gray to black lateral stripe, and a larger, slightly oblique mouth that extends to or slightly beyond the anterior edge of the eye. Southern Redbelly Dace have a moderately oblique, slightly subterminal mouth with the upper jaw slightly protruding past the lower jaw, and a less blunt snout.

Northern Pearl Dace display one thick, dark lateral stripe outlined by two thin, faint golden bronze lateral stripes, have one barbel at each corner of the mouth that are sometimes present, have larger scales, and a complete lateral line.

Distribution and Habitat: Native across Canada and the northern United States from

Montana in the west, east throughout northern Missouri, Mississippi, Great Lakes-St.

Lawrence and Atlantic river drainages to Maine in the east, and south to Nebraska with isolated populations in Colorado. Within South Dakota, Northern Redbelly Dace are found primarily east of the Missouri River and have been reported from the tributaries of the Missouri, Big Sioux, Minnesota, White, Niobrara, and Keya Paha river drainages, which are on the southern periphery of the geographic range for Northern Redbelly

Dace.1,6,9,10,13 Northern Redbelly Dace have been reported from the upper reaches of the

Knife, Heart, and Cannonball river drainages in western North Dakota, in Spring Creek and Clear Creek in central North Dakota, and the Red River of the North drainage in eastern North Dakota.13,15,17 Well adapted to cold, clear waters of first order springfed streams in areas associated with vegetation, low velocity, and sand or gravel substrate.18

Exhibits natal site fidelity.12 Often found within the littoral zones of lakes in schools of

50 to roughly 300 individuals.8 In Canadian Shield lakes, Northern Redbelly Dace have been reported to exhibit onshore-offshore diel migrations, occupying the littoral zones 158 during the day, the pelagic zones in the evenings, and migrating back to the littoral zone during sunrise.14 In littoral zones, the species has shown preference for densely covered or structured habitat over open water.7,14 Northern Redbelly Dace likely inhabit the littoral zone during the day to avoid predation by pelagic piscivores.7,14 The species has demonstrated fright reactions to skin extracts or chemicals of conspecifics by increasing dashing and freezing behaviors, moving towards the substrate and fleeing the area where the alarm substance is present, and increasing the cohesiveness of schools.8 Individuals show stronger and longer fright reactions and behaviors when concentrations of alarm substances are increased, suggesting the ability of the species to detect different degrees of predation risk.8 These chemical cues and antipredator responses may increase survival rates, especially when visual cues are limited.8

Reproduction: Little is known about the Northern Redbelly Dace’s reproductive behavior, but it is believed to spawn during late spring to early summer.2 As many as four to five males have been observed following a single female into a mass of filamentous algae, where they will push up against her and vibrate their bodies to initiate the release of eggs and milt.3,5 Sexual maturity reached at age 1-2.2 Fractional spawners, with roughly 5-30 eggs released during a single spawning event.5 No nest is constructed and no parental care is given. Eggs yellow-orange in color, roughly 0.9 mm (0.04 in) in diameter, and nonadhesive.2 Hatching takes place within 8-10 days in water 21.1-26.7°C

(69.98-80.06°F).2,11

Age and Growth: Information on age and growth is not well understood. Capable of reaching 77 mm (3.03 in) TL.2 Longevity is 8 years.16 159

Food and Feeding: Adults primarily feed within the water column on large amounts of green algae, detritus and small, immature aquatic invertebrates.4 Visual omnivores, diel migration movements to the pelagic zone in the evenings is likely to increase the species feeding efficiency on zooplankton.8

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Miscellaneous Publication No. 119. Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Bertrand-Toline, C.A. 1994. Morphometric and genetic differentiation of the

northern redbelly dace: Investigating microevolutionary processes. Ph.D.

Dissertation, University of Toronto.

4. Cochran, P.A., D.M. Lodge, J.R. Hodgson, and P.G. Knapik. 1988. Diets of

syntopic finescale dace, Phoxinus neogaeus, and northern redbelly dace, Phoxinus

eos: a reflection of trophic morphology. Environmental Biology of Fishes 22:235-

240.

5. Cooper, G.P. 1935. Some results of the forage fish investigations in Michigan.

Transactions of the American Fisheries Society 65:132-142.

6. Dieterman, D., and C.R. Berry Jr. 1994. Fishes in seven streams of the Minnesota

River drainage in northeastern South Dakota. Proceedings of the South Dakota

Academy of Science 73:23-30.

7. Dupuch, A., P. Magnan, A. Bertolo, L.M. Dill, and M. Proulx. 2009. Does

predation risk influence habitat use by northern redbelly dace Phoxinus eos at

different spatial scales? Journal of Fish Biology 74:1371-1382. 160

8. Dupuch, A., P. Magnan, and L.M. Dill. 2004. Sensitivity of northern redbelly

dace, Phoxinus eos, to chemical alarm cues. Canadian Journal of Zoology 82:407-

414.

9. Felts, E.A. 2013. Ecology of glacial relict fishes in South Dakota’s sandhills

region. M.S. Thesis, South Dakota State University, Brookings.

10. Felts, E.A., and K.N. Bertrand. 2014. Conservation status of five headwater

stream specialists in South Dakota. The American Midland Naturalist 172:131-

159.

11. Hubbs, C.L., and G.P. Cooper. 1936. Minnows of Michigan. Cranbrook institute

Science Bulletin 8:1-95.

12. Massicotte, R., P. Magnan, and B. Angers. 2008. Intralacustrine site fidelity and

nonrandom mating in the littoral-spawning northern redbelly dace (Phoxinus eos).

Canadian Journal of Fisheries and Aquatic Sciences 65:2016-2025.

13. Morey, N.M., and C.R. Berry Jr. 2004. New distributional record of the Northern

Redbelly Dace in the Northern Great Plains. The Prairie Naturalist 36:257-260.

14. Naud, M., and P. Magnan. 1988. Diel onshore-offshore migrations in northern

redbelly dace, Phoxinus eos (Cope), in relation to prey distribution in a small

oligotrophic lake. Canadian Journal of Zoology 66:1249-1253.

15. Reigh, R.C., and J.B. Owen. 1978. Fishes of the western tributaries of the

Missouri River in North Dakota. North Dakota Regional Environmental

Assessment Program, Bismarck, North Dakota.

16. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p. 161

17. Stasiak, R.H. 1980. Phoxinus eos (Cope), northern redbelly dace. Pages 336 in

Atlas of North American freshwater fishes (D.S. Lee, C.R. Gilbert, C.H. Hocutt,

R.E. Jenkins, D.E. McAllister, and J.R. Stauffer, Jr., editors) North Carolina State

Museum of Natural History, Raleigh, North Carolina.

18. Stasiak, R.H. 2006. Northern redbelly dace (Phoxinus eos): A technical

conservation assessment. USDA Forest Service, Rocky Mountain Region. 42p.

162

Southern Redbelly Dace, Chrosomus erythrogaster (Rafinesque, 1820)

Etymology and Synonyms: Chrosomus = “coloration”; erythro = “red”, gaster =

“belly”, referring to the red-orange flank coloration of the species, especially on males.

Description: Body fusiform, elongate, slightly laterally compressed. Dorsally olive to gray with small irregular, dark specks; lateral sides with multiple stripes of color, beginning with a thin, narrow dark lateral stripe nearest to dorsal side (beginning at tip of snout and extending to base of caudal fin), then a thicker silvery white to cream lateral stripe below, followed by a thicker dark black lateral stripe (beginning at tip of snout and extending to base of caudal fin creating a dark caudal spot), and males display light, dusky red to orange flanks below the thicker, dark lateral stripe; ventrally cream to white; fins generally clear but may display light yellow pigmentation. Head moderate. Snout moderately pointed. Eye moderately large, placed laterally on head. Mouth small, moderately oblique and slightly subterminal; upper jaw protruding past lower jaw; jaws do not extend past anterior end of eye. Lips fleshy. Barbels absent. Pharyngeal teeth slender; pattern 0,5-5,0. Gill rakers 7-10, short. Dorsal fin with 8 rays. Adipose fin absent. Caudal peduncle elongate, thick. Caudal fin moderately forked. Anal fin with 8 rays. Pelvic fins abdominal with 8 rays; insertion distinctly anterior to insertion of dorsal fin. Pectoral fins short, rounded with 14-17 rays. Lateral line incomplete with 70-95 small cycloid scales in series. Spawning males develop intense, bright nuptial coloration including bright red-orange on flanks below lateral stripe, and orange-yellow coloration at base of dorsal, anal, pelvic and pectoral fins. Spawning females also develop nuptial coloration, but to a much lesser extent and appearing very faint. Juveniles similar to adults with less coloration. 163

Similar Species: Closely resembles Northern Redbelly Dace, Northern Pearl Dace, and

Finescale Dace. Northern Redbelly Dace have a blunt snout, a terminal and strongly oblique mouth, and a lower jaw that often protrudes past the upper jaw. Northern Pearl

Dace display one thick, dark lateral stripe outlined by two thin, faint golden bronze lateral stripes, have one barbel at each corner of the mouth that are sometimes present, have larger scales, and a complete lateral line. Finescale Dace display a single distinct dark gray to black lateral stripe, and a larger, slightly oblique mouth that extends to or slightly beyond the anterior edge of the eye.

Distribution and Habitat: Widely distributed throughout the upper Mississippi, Ohio and Missouri River drainages in the United States from southern Minnesota and

Wisconsin in the north, east to Pennsylvania and New York, south to the Ozark regions of Arkansas and Oklahoma, as well as the lower Tennessee River drainage, and to New

Mexico and Colorado in the west.10,14,15 Absent from North Dakota and have only been reported in eastern South Dakota within tributaries of the Big Sioux River in Lincoln

County.12,15 Inhabits small to medium low-order streams in waters generally 0.1-1.5 m

(0.33-4.92 ft) in depth.1,10,15 Abundant and often found schooling in small spring-fed headwater reaches and streambeds with cool, clear waters within slow-flowing pools

(0.05-0.20 m/ sec) below shallow riffles and runs with detritus, gravel, pebble, sand and small boulder substrate and dense riparian canopy cover.7,10 Also prefers the cover of undercut banks and areas with permanent vegetation.15 Avoids areas with greater amounts of clay, silt, mud, and large boulder substrate.1,10 Critical thermal maxima with an endpoint of death recorded at 35 °C (95 °F).4 164

Reproduction: Spawning takes place May to June, with older adults spawning earlier in the season and younger adults spawning later in the season.7,9,11 Spawning ceases when water temperature reaches 21 °C (69.8 °F).5,9 Individuals <40 mm (1.57 in) TL are considered immature, with most reaching sexual maturity at age-1, or approximately 50 mm (1.97 in) TL.15 Spawning occurs over gravel, pebble or sand substrate within swift flowing waters.5,11 No nest is constructed, but the species is often observed spawning over a nest constructed by other cyprinid species including Creek Chub, stonerollers and

Common Shiners.3,5,7 Males temporarily defend small territories around a female.5

Spawning behavior begins with several males pursuing a single female, and later two males nudging and crowding laterally against a single female, clasping onto her by placing one pectoral fin under her body and curving his body close to hers.5,6,7,11 The males body vibrates rapidly against the females, initiating the release of eggs and milt.11

Broadcast spawners.13 Average fecundity for ages 0-2 were 267, 401, and 568 respectively.9,15 Eggs roughly 0.75-1.3 mm (0.03-0.05 in) in diameter and nearly transparent.1,11 Hatching occurs within six days at a water temperature of 20 °C (68

°F).15,16

Age and Growth: Females generally smaller than males the same age.1,15 Capable of reaching 88.9 mm (3.5 in) TL.3 Longevity 4 years.15

Food and Feeding: Adults are generalist, visual, omnivorous grazers who feed on algae and aquatic invertebrates.2,8 Moderate densities of Southern Redbelly Dace can temporarily reduce mean algal filament length and mean size of particulate organic matter.2

Literature Cited: 165

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Bertrand, K.N., and K.B. Gido. 2007. Effects of the herbivorous minnow,

southern redbelly dace (Phoxinus erythrogaster), on stream productivity and

ecosystem structure. Oecologia 151:69-81.

3. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

4. Farless, N.A., and S.K. Brewer. 2017. Thermal tolerance of fishes occupying

groundwater and surface-water dominated streams. Freshwater Science 36:866-

876.

5. Hayes, B.M. 2015. Reproductive biology of syntopic blackside dace and southern

redbelly dace in two Kentucky streams. M.S. Thesis, Eastern Kentucky

University, Richmond.

6. Johnston, C., and L. Page. 1992. The evolution of complex reproductive strategies

in North American minnows (Cyprinidae). Pages 600-621 in R. Mayden (editor).

Systematics, historical ecology, and North American freshwater fishes. Stanford

University Press, Stanford, California.

7. Phillips, G.L. 1968. Chrosomus erythrogaster and Chrosomus eos (Osteichthyes:

Cyprinidae): taxonomy, distribution, ecology. Ph.D. Dissertation, Department of

Zoology, University of Minnesota.

8. Settles, W.H., and R.D. Hoyt. 1976. Age structure, growth patterns, and food

habits of the southern redbelly dace Chrosomus erythrogaster in Kentucky.

Transactions of the Kentucky Academy of Science 37:1-10. 166

9. Settles, W.H., and R.D. Hoyt. 1978. The reproductive biology of the southern

redbelly dace, Chrosomus erythrogaster Rafinesque, in a spring-fed stream in

Kentucky. American Midland Naturalist 99:290-298.

10. Slack, W.T., M.T. O’Connell, T.L. Peterson, J.A. Ewing III, and S.T. Ross. 1997.

Ichthyofaunal and habitat associations of disjunct populations of southern

redbelly dace. Phoxinus erythrogaster (Teleostei: Cyprinidae) in Mississippi. The

American Midland Naturalist 137:251-265.

11. Smith, B.G. 1908. The spawning habits of Chrosomus erythrogaster Rafinesque.

The Biological Bulletin 15:9-18.

12. Springman, D.J., and R.L. Banks. 2005. Range extension of the southern redbelly

dace into South Dakota. The Prairie Naturalist 37:175-176.

13. Starnes, L.B., and W.S. Starnes. 1981. Biology of the blackside dace Phoxinus

cumberlandensis. American Midland Naturalist 106:360-371.

14. Starnes, W.C., and L.B. Starnes. 1980. Phoxinus erythrogaster (Rafinesque),

southern redbelly dace. Page 337 in D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E.

Jenkins, D.E. McAllister, and J.R. Stauffer, Jr. (editors). Atlas of North American

freshwater fishes. North Carolina State Museum of Natural History, Raleigh.

15. Stasiak, R.H. 2007. Southern redbelly dace (Phoxinus erythrogaster): a technical

conservation assessment. U.S. Department of Agriculture, Forest Service, Rocky

Mountain Region, Golden, Colorado.

16. Sternburg, J.G. 1992. Spawning of southern and northern redbelly dace. North

American Native Fishes Association, winter 1992-1993.

167

Finescale Dace, Chrosomus neogaeus (Cope, 1867)

Etymology and Synonyms: Chrosomus = “coloration”; neogaeus = new world.

Description: Body fusiform, anteriorly cylindrical, laterally compressed posteriorly.

Dorsally dusky black to brown; laterally light gray to olive followed by a single distinct dark gray to black lateral stripe extending from tip of snout to base of caudal fin, sometimes leading to a dark caudal spot; ventrally light gray fading to white on midventral surface; fins lightly pigmented. Head large. Snout blunt. Eye moderately large, placed laterally on head. Mouth large, terminal, slightly oblique; upper jaw extends to anterior end or middle of eye. Lips fleshy, may slightly protrude past tip of snout.

Barbels absent. Pharyngeal teeth robust and hooked; pattern 2,5-4,2. Gill rakers short, 9.

Dorsal fin with rounded distal end and 8 rays. Adipose fin absent. Caudal peduncle elongate, uniform in thickness. Caudal fin moderately forked. Anal fin with rounded distal end and 8 rays. Pelvic fins abdominal with rounded distal end and 8 rays; insertion distinctly anterior to insertion of dorsal fin. Pectoral fins with rounded distal ends and 15-

16 rays. Lateral line incomplete and short with 70-85 extremely small ctenoid scales in series, appearing scaleless with the naked eye. Spawning males develop a distinct burnt orange to red lateral stripe immediately below black lateral stripe; females with a dusky, faded burnt orange to yellow lateral stripe. Males also develop thickened, highly modified pectoral fin rays and rows of tubercles on the ventral region, above base of the anal fin, and near the opercle.

Similar Species: Closely resembles Northern Redbelly Dace. Northern Redbelly Dace have two dark black to brown lateral stripes separated by a thicker light olive to gray stripe, and a small mouth with an upper jaw that does not extend to or beyond anterior 168 end of eye. Southern Redbelly Dace have a more elongate snout, small dark spots on dorsal side and two dark black to brown lateral stripes separated by a thicker light gray to white stripe. Northern Pearl Dace may have one barbel at each corner of the mouth that are sometimes present, have larger scales, and a complete lateral line.

Distribution and Habitat: Native across the glaciated areas of southern Canada and the northern United States from northern Montana in the west, south to disjunct populations in Wyoming, Nebraska and South Dakota within the upper Missouri River basin, and north into the Mississippi and Great Lakes-St. Lawrence river drainages to Maine.2,7 In

North Dakota Finescale Dace occur within the Red River tributaries and along the

Missouri River, and in southwestern South Dakota have been recorded in the Belle

Fourche, Cheyenne, White, and Niobrara watersheds.1,2,6 In the northern portion of the species range, Finescale Dace are common in springfed streams with low velocity, and are often associated with beaver ponds.6,8,9,11 Most frequent in cold water first order streams.6 Also inhabit glacial lakes, ponds and bogs with cool water and abundant cover.9,10 Strongly associated with shorelines and structure.4 Intolerant of water temperature >25 °C (77 °F).7

Reproduction: Spawning takes place late April through late May, and may extend into

June or early July depending on water temperature.5,10 In Minnesota, spawning took place when water temperatures began to exceed 15 °C (59 °F), and continued until 22 °C (71.6

°F) in shallow areas 0.5-0.9 m (1.64-2.95 ft) deep.3,10 Sudden fluctuations in water temperature caused by ice-out are suggested to initiate spawning.9 Sexual maturity reached at age 1-2.10 Spawning behavior consists of one or two ripe females leaving a school to take cover often in overhanging vegetation or woody debris, where they are 169 followed by several males.10 A male will grasp onto a female by placing his thickened, modified pectoral fins on the females ventral side just posterior to her pectoral fins.10 He then forces the female against a sturdy object and curves his caudal fin over the females caudal fin, causing the patch of tubercles above the base of his anal fin to rub the vent of the female.10 This motion initiates the release of milt and roughly 20-30 eggs.10 Roughly ten seconds after the spawning action ceases, females retreat back into schools and males linger for several more seconds continuing releasing milt.10 No nest is constructed and no parental care is given.10 Fractional spawners. Fecundity rather low for cyprinid species, but increases with length of female, and ranges 784-3,060 eggs per female.10 Eggs yellow-orange in color, roughly 1.24-1.5 mm (0.05-0.06 in) in diameter, and demersal.5,9,10 Hatching occurs in 6 days in water 20 °C (68 °F).10

Age and Growth: Newly hatched larvae roughly 4.2 mm (0.17 in) TL.10 Standard length of both sexes roughly equal after first year of growth.10 Females tend to live longer and have faster growth rates than males following sexual maturity.10 Capable of reaching 107 mm (4.21 in) TL.3 Longevity 6 years.10

Food and Feeding: Opportunistic omnivore. Adults feed within shallow shoreline areas, and diets are composed of macroinvertebrates such as larval Chironomids, immature

Odonata, Trichoptera larvae, Ephemeropterans and Coleopterans.4 Adults also consume small mollusks, vegetative matter, green algae, diatoms and zooplankton.4

Literature Cited:

1. Amiotte, J., G. Simpson, and M.E. Barnes. 2015. Re-establishment of finescale

dace (Phoxinus neogaeus) in Mud Lake, Lawrence County, South Dakota.

Proceedings of the South Dakota Academy of Science 94: 195-200. 170

2. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Cochran, P.A., D.M. Lodge, J.R. Hodgson, and P.G. Knapik. 1988. Diets of

syntopic finescale dace, Phoxinus neogaeus, and northern redbelly dace, Phoxinus

eos: a reflection of trophic morphology. Environmental Biology of Fishes 22:235-

240.

5. Das, M.K., and J.S. Nelson. 1990. Spawning time and fecundity of northern

redbelly dace, Phoxinus eos, finescale dace, Phoxinus neogaeus, and their hybrids

in Upper Pierre Lake, Alberta. Canadian Field-Naturalist 104:409-413.

6. Felts, E.A., and K.N. Bertrand. 2014. Conservation status of five headwater

stream specialists in South Dakota. The American Midland Naturalist 172:131-

159.

7. Isaak, D.J., W.A. Hubert, and C.R. Berry. 2003. Conservation assessment of the

lake chub, mountain sucker, and finescale dace in the Black Hills National Forest,

South Dakota and Wyoming. USDA Forest Service, Rocky Mountain Region.

8. Scholsser, I.J. 1995. Dispersal, boundary processes, and trophic-level interactions

in streams adjacent to beaver ponds. Ecology 76:908-925.

9. Stasiak, R.H. 1972. The morphology and life history of the finescale dace, Pfrille

neogaea, in Itasca State Park, Minnesota. Ph.D. dissertation. University of

Minnesota, Minneapolis. 171

10. Stasiak, R.H. 1978. Reproduction, age, and growth of the finescale dace,

Chrosomus neogaeus, in Minnesota. Transactions of the American Fisheries

Society 107:720-723.

11. Stasiak, R.H., and G.R. Cunningham. 2006. Finescale dace (Phoxinus neogaeus):

a technical conservation assessment. USDA Forest Service, Rocky Mountain

Region. 49p.

172

Lake Chub, Couesius plumbeus (Agassiz, 1850)

Etymology and Synonyms: Couesius = named after Dr. Elliot Coues, American historian and ornithologist; plumbeus = lead colored, possibly referring to the dorsal coloration or lateral stripe.

Description: Body fusiform, elongate, rather cylindrical anteriorly and moderately compressed posteriorly. Dorsally blueish gray to brown; laterally light gray to silver with faint mottling and dusky, faint lead colored lateral stripe; ventrally silver to white; fins generally clear to light gray without any distinctive markings. Head small, moderately dorsoventrally compressed. Snout bluntly pointed, slightly extending beyond the mouth.

Eye moderately large, placed laterally on upper portion of head. Mouth relatively small with upper jaw barely reaching to anterior end of eye. Barbels present, one on each corner of the mouth; moderately long, prominent. Lips thin. Pharyngeal tooth pattern 2,4-

4,2. Gill rakers short, 4-8. Dorsal fin with 8 rays, nearly straight to slightly concave distal end with rounded tip. Adipose fin absent. Caudal peduncle moderately thick. Caudal fin forked. Anal fin with 8 rays and straight distal end. Pelvic fins abdominal with 8 rays; insertions anterior to insertion of dorsal fin. Pectoral fins short with 15-17 rays; do not extend beyond insertions of pelvic fins when depressed. Lateral line complete with 53-70 small cycloid scales in series. Spawning males develop nuptial coloration including red to orange coloring on the bases of the paired fins, red coloration near the mouth and opercle, and a more pronounced lateral stripe. Spawning females also develop nuptial coloration including faint red coloring on the bases of the pectoral fins. Both sexes develop small breeding tubercles on the head, cheeks, dorsal side of the body, and the pectoral and pelvic fins. 173

Similar Species: Closely resembles the Creek Chub. Creek Chub have a large mouth with an upper jaw that extends past the anterior edge of the eye and have a large dark spot present on the anterior, basal portion of the dorsal fin.

Distribution and Habitat: Native to North America from the Atlantic basin in Canada and the United States, west to the Rocky Mountains, north and west to the Pacific basin in British Columbia, and north and east to the Arctic basin in northwestern Canada,

Alaska and Labrador.4,9 Primarily occurs throughout the upper Missouri, Little Missouri, and Knife River watersheds in North Dakota. Considered rare in Lake Sakakawea and

Fort Peck Reservior.11 Lake Chub distribution in South Dakota is currently declining, however the species is still present in the Little Missouri and Cheyenne River drainages.7,8 The decline of the species within South Dakota may be a result of water withdrawals and warming water temperatures in streams due to siltation and the destruction of riparian zones.7 Introduced trout may also be negatively impacting Lake

Chub populations via predation and competition.7 Inhabit cool, clear waters of lakes and small streams within environments that experience great seasonal variations in water temperature, ranging approximately 4 °C (39.2 °F) in the winter to the mid-20 °C (68 °F) in summer.4,6,10 Often occurs in large schools within shallow waters over sand or gravel substrate.1 Tolerant of moderate turbidity.1,10

Reproduction: Spawning takes place when water temperatures reach roughly 10 °C (50

°F), which may occur throughout the spring or summer depending on location.1,2,3

Migration movements upstream to spawning grounds may be as large as 3 km (1.86 mi).3

Sexual maturity occurs at age-3, or when individuals reach approximately 95.1mm (3.74 in) TL.2,3 Males known to display some aggressive behavior during spawning, but are not 174 territorial.3 Males court females within the spawning grounds by a combination of nudging and persistent pursuit movements.3 Spawning behavior consists of a single male pressing a single female against the substrate or a rock, and vibrating his body against hers to initiate the release of eggs and milt.2 This spawning act lasts roughly 1 second, and is known to occur several times between the same pair of individuals, with a few eggs being released at a time.2 Broadcast spawner, with eggs being dispersed over silt, gravel, or rocky substrate.1,3 No nest is constructed and no parental care is given.3

Fecundity generally increases with the size of female.1,2 Fecundity of females 168-

181mm (6.61-7.13 in) TL ranged 5,290-6,630 eggs per female.1 Eggs light yellow in color, roughly 1.6-3.16 mm (0.06-0.13 in) in diameter and nonadhesive.1,2,3 Hatching takes place in roughly 10 days at a water temperature of 13.33 °C (56 °F).2

Age and Growth: Information on age and growth of the species is scarce, especially in the Dakotas. Average lengths-at-age of individuals from Montana are reported as: age-1,

63.5 mm (2.5 in) TL; age-2, 114.3 mm (4.5 in) TL; age-3, 139.7 mm (5.5 in) TL, which are similar to average lengths-at-age reported from Wisconsin.1,2 Capable of reaching 227 mm (8.94 in) TL.12 Longevity 7 years, with few individuals surviving beyond age-5.1

Food and Feeding: Information regarding the diet and feeding habits of Lake Chub is limited. Lake Chub are likely visual feeders given their lack of external taste buds on the body.5 Juvenile diets primarily consist of small microcrustaceans such as copepods and cladocerans.1 Adults primarily forage on small aquatic invertebrates such as ephemeroptera, diptera, and odonatan larvae.1 Lake Chub have also been known to consume plankton and small amounts of crustaceans.1

Literature Cited: 175

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

3. Brown, J.H., U.T. Hammer, and G.D. Koshinsky. 1970. Breeding biology of the

Lake Chub, Couesius plumbeus, at Lac la Ronge, Saskatchewan. Journal of the

Fisheries Research Board of Canada 27:1005-1015.

4. Darveau, C.A., E.B. Taylor, and P.M. Schulte. 2012. Thermal physiology of

warm-spring colonists: variation among lake chub (Cyprinidae: Couesius

plumbeus) populations. Physiological and Biochemical Zoology 85:607-617.

5. Davis, B.J., and R.J. Miller. 1967. Brain patterns in minnows of the genus

Hybopsis in relation to feeding habits and habitat. Copeia 1967:1-39.

6. Evermann, B.W., and U.O. Cox. 1896. Report upon the fishes of the Missouri

River basin. In Report of the U.S. Commissioner of fish and fisheries for 1894.

U.S. Government Printing Office, Washington, D.C.

7. Hoagstrom C.W., C.A. Hayer, J.G. Kral, S.S. Wall, and C.R. Berry Jr. 2006. Rare

and declining fishes of South Dakota: a river drainage scale perspective.

Proceedings of the South Dakota Academy of Science 85:171-211.

8. Isaak, D.J. W.A. Hubert, and C.R. Berry, Jr. 2003. Conservation assessment for

lake chub, mountain sucker, and finescale dace in the Black Hills National Forest,

South Dakota and Wyoming. U.S. Department of Agriculture, Forest Service,

Custer, South Dakota. 176

9. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

10. McPhail, J.D., and C.C. Lindsey. 1970. Freshwater fishes of northwestern Canada

and Alaska. Bulletin 173. Fisheries Research Board of Canada, Ottawa.

11. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

12. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research. Board of Canada, Bulletin 184. 966p.

177

Grass Carp, Ctenopharyngodon idella (Valenciennes, 1844)

Etymology and Synonyms: Cteno = Greek for “comb”, pharyn = Greek for “throat”, odon = Greek for “tooth”, referring to the comb-like and sharp pharyngeal teeth; idella = derived from the Greek word “ideo” meaning distinct.

Description: Body fusiform, slightly elongate, rather cylindrical. Dorsally olive to gray in color; laterally light olive to silver or bronze with scales outlined in darker color appearing crosshatched; ventrally silver to white; fins dark olive to gray in color. Head broad and small. Snout slightly conical, short. Eye moderately large, placed laterally in middle of head. Mouth large, terminal; upper jaw barely extends backward towards anterior edge of eye. Barbels absent. Lips thin. Teeth on jaws absent. Pharyngeal teeth deeply grooved, sharp, comb-like; 2,4-5,2 or 2,5-4,2. Gill rakers short, unfused; 15-16.

Dorsal fin short with 7-9 rays and no spine. Adipose fin absent. Caudal peduncle slightly elongate, thick. Caudal fin forked. Anal fin set far posterior on body with 8-10 rays and no spine. Pelvic fins abdominal with 7-8 rays and no spine; insertion slightly posterior to insertion of dorsal fin. Ventral keel absent. Pectoral fins with 18-20 rays. Lateral line complete, slightly decurved with 34-43 large cycloid scales in series. Spawning males with minute tubercles on pectoral fins. Juveniles similar to adults, and may be more silver in color.

Similar Species: Easily distinguished from Common Carp by the elongate and cylindrical body shape, and lack of barbels. Bighead and Silver Carp have much smaller cycloid scales and are deeper and laterally compressed. Buffalo (Ictiobus) species are less elongate and have a much more elongate dorsal fin. 178

Distribution and Habitat: Native to large rivers and lakes of southern Russia to northern China and Vietnam, but have been extensively introduced throughout the world.7 Imported to the United States in 1963 as means of aquatic vegetation control, and first accidentally released in 1966 by the U.S. Fish and Wildlife Service in Arkansas from aquaculture facilities.10,16 In the United States, the species is now established in the

Mississippi River basin, as well as the Trinity River basin in Texas.4 Found east of the

Missouri River in the Dakotas. Inhabit shallow, densely vegetated areas of reservoirs, lakes, ponds, backwaters or pools of large rivers, and occasionally small streams.14,18

Most active during daylight hours.14 Negatively impact aquatic ecosystems by overgrazing native macrophytes. Tolerant of a wide range of water temperatures from 0-

33 °C (32-91.4 °F), as well as rapid changes in temperature.7,8 Mean critical thermal maximum 39.3 °C (102.74 °F).5,7 Lower lethal temperature tolerance of fry is 0-0.1 °C

(32-32.18 °F), and upper lethal water temperature is 33-41 °C (91.4-105.8 °F).5,7 Able to tolerate oxygen concentrations as low as 0.2 mg/L.7,18

Reproduction: Opportunistic spawners.20 Spawning takes place during spring and early summer, or when water temperatures reach 20-30 °C (68-86 °F) in turbid and turbulent waters of main channels in rivers and canals during high water.7,8,18 Optimum spawning temperature 20-22 °C (68-71.6 °F), but known to spawn at temperatures as low as 15 °C

(59 °F).7,18 Sexual maturity occurs at age 4-5 in temperate areas of the United States.7

Males mature roughly one year prior to females.7 One female is accompanied by two or more males during the spawning act.7 Females highly fecund, averaging 1 million eggs per female.1,20 No nest is constructed and no parental care is given. Eggs roughly 2.0-2.5 mm (0.08-0.10 in) in diameter, semibuoyant and non-adhesive.7 Optimum incubation 179 temperature 21-26 °C (69.8-78.8 °F).7,8,18 Hatching occurs within 26-60 hours at 17-30

°C (62.6-86 °F).7 Triploid Grass Carp incapable of reproducing are often stocked as biological control for aquatic weeds.2,10

Age and Growth: Growth rapid during early stages. In hatchery ponds, juvenile grass carp growth rates were 9.8 g/day (0.002 lbs/day).12 Under optimal conditions, Grass Carp often reach 1.0 kg (2.20 lbs) during their first year.22 In a Florida lake, 90 g (0.20 lbs)

Grass Carp experienced a growth rate of 10.0 g/day (0.02 lbs/day) one year post stocking.19 The growth rate of yearling Grass Carp and Bighead Carp hybrids in South

Dakota was roughly one-third of similar sized Grass Carp.9 Capable of reaching >1000 mm (39.37 in) TL, and generally attains weights of 30-50 kg (66.14-110.23 lbs).7

Lifespan may be longer at higher latitudes with lower water temperatures and shorter growing seasons.11 Longevity 15 years, although one specimen from North Dakota was found to be over 33 years old.2,7,11

Food and Feeding: Adults largely opportunistic herbivores, and feed heavily upon submerged rooted aquatic macrophytes, as well as benthic invertebrates, filamentous algae, and sometimes flooded terrestrial vegetation.6,18 Majority of active feeding takes place during the morning and evening, with maximum consumption occurring at 22-23

°C (71.6-73.4 °F).15,17 During this optimum feeding temperature, Grass Carp are known to consume 110-169 % of their body weight/day.3,15 In South Dakota, Grass Carp and

Bighead Carp hybrids primarily feed diurnally in shallow waters, and heavily consumed

Najas guadalupensis and Chara sp., as well as Potamogeton pectinatus.9 The species feeding habits often times have serious detrimental effects on biotic and abiotic factors of aquatic ecosystems and their native fish species and even waterfowl.7,13 Larvae primarily 180 feed on zooplankton such as rotifers.21 Hybrids consume roughly one-third of what Grass

Carp consume daily.9

Literature Cited:

1. Aliev, D.S., and A.I. Sukhanova. 1974. Fecundity of the grass carp

Ctenopharyngodon idella (Val.) and the silver carp Hypophthalmichthys molitrix

(Val.) in the Kara-kum canal and its reservoirs. Izv. Akad. Nauk Turkm. SSR Ser.

Biol. Nauk 4:77-83.

2. Allen, S.K., and R.J. Wattendorf. 1987. Triploid grass carp: status and

management implications. Fisheries 12:20-24.

3. Caldwell, B.A. 1980. Ability of grass carp to control aquatic macrophytes in fish

culture ponds in Colorado. M.S. Thesis, Colorado State University, Fort Collins.

4. Chapman, D.C., and M.H. Hoff. 2011. Introduction. Pages 1-4 in D.C. Chapman

and M.H. Hoff, editors. Invasive Asian carps in North America. American

Fisheries Society, Symposium 74, Bethesda, Maryland.

5. Chilton III, E.W., and M.I. Muoneke. 1992. Biology and management of grass

carp (Ctenopharyngodon idella, Cyprinidae) for vegetation control: a North

American perspective. Reviews in Fish Biology and Fisheries 2:283-320.

6. Colle, D.V., J.V. Shireman, and R.W. Rottman. 1978. Food selection by grass

carp fingerlings in a vegetated pond. Transactions of the American Fisheries

Society 107:149-152.

7. Cudmore, B., and N.E. Mandrak. 2004. Biological synopsis of grass carp

(Ctenopharyngodon idella). Canadian Manuscript Report of Fisheries and

Aquatic Sciences 2705. 181

8. Fedorenko, A.Y., and F.J. Fraser. 1978. Review of grass carp biology.

Interagency Committee on Transplants and Introductions of Fish and Aquatic

Invertebrates in British Columbia, Department of Fisheries and Environment,

Fisheries and Marine Service, Technical Report No. 786.

9. Harberg, M.C., and T. Modde. 1985. Feeding behavior, food consumption,

growth, and survival of hybrid grass carp in two South Dakota ponds. North

American Journal of Fisheries Management 5:457-464.

10. Kelley, A.M., C.R. Engle, M.L. Armstrong, M. Freeze, and A.J. Mitchell. 2011.

History of introduction and governmental involvement in promoting the use of

grass, silver, and bighead carps. Pages 163-174 in D.C. Chapman and M.H. Hoff,

editors. Invasive Asian carps in North America. American Fisheries Society,

Symposium 74, Bethesda, Maryland.

11. Kirk, J.P., and R.C. Socha. 2003. Longevity and persistence of triploid grass carp

stocked into the same Santee Cooper reservoirs of South Carolina. Journal of

Aquatic Plant Management 41:90-92.

12. Lembi, C.A., B.G. Ritenuor, E.M. Iverson, and E.C. Forss. 1978. The effects of

vegetation removal by grass carp on water chemistry and phytoplankton in

Indiana ponds. Transactions of the American Fisheries Society 107:161-171.

13. McKnight, S.K., and G.R. Hepp. 1995. Potential effect of grass carp herbivory on

waterfowl foods. Journal of Wildlife Management 59:720-727.

14. Nixon, D.E., and R.L. Miller. 1978. Movements of grass carp, Ctenopharyngodon

idella, in an open reservoir system as determined by underwater telemetry.

Transactions of the American Fisheries Society 107:146-148. 182

15. Opuszynski, K. 1972. Use of phytophagous fish to control aquatic plants.

Aquaculture 1:61-74.

16. Pflieger, W.L. 1978. Distribution and status of the grass carp (Ctenopharyngodon

idella) in Missouri streams. Transactions of the American Fisheries Society

107:113-118.

17. Shireman, J.V., and C.R. Smith. 1981. Biological synopsis of grass carp

(Ctenopharyngodon idella). Final Report, contract number 14-16-0009-78-912.

U.S. Fish and Wildlife Service, Washington, D.C.

18. Shireman, J.V., and C.R. Smith. 1983. Synopsis of biological data on the grass

carp Ctenopharyngodon idella (Cuvier and Valenciennes, 1844). FAO Fisheries

Synopsis 135.

19. Shireman, J.V., D.E. Colle, and M.J. Maceina. 1980. Grass carp growth rates in

Lake Wales, Florida. Aquaculture 19:379-382.

20. Stanley, J.G. 1976. Reproduction of the grass carp (Ctenopharyngodon idella)

outside its native range. Fisheries 1:7-10.

21. Stanley, J.G., W.W. Miley II, and D.L. Sutton. 1978. Reproductive requirements

and likelihood for naturalization of escaped grass carp in the United States.

Transactions of the American Fisheries Society 107:119-128.

22. Vietmeyer, N.D. 1976. Grass carp. Pages 15-21 in N.D. Vietmeyer, editor.

Making aquatic weeds useful: some perspectives for developing countries.

National Academy of Sciences, Washington, D.C., USA.

183

Red Shiner, Cyprinella lutrensis (Baird & Girard, 1853)

Etymology and Synonyms: Cyprinella = Latin for “cyprinus”, meaning “carp”; lutrensis = lutra, referring to Otter; the species was originally documented in Otter Creek,

Arkansas.1

Description: Body fusiform, deep and laterally compressed. Dorsally light olive green to dusky gray with scales outlined in darker pigment creating a diamond shaped pattern; laterally silvery blue with scales outlined in a darker pigment creating a diamond shaped pattern; ventrally silver to white; dorsal fin clear to light gray; caudal, anal and paired fins clear to faint reddish-orange. Head short, deep. Snout short, tapering to a blunt point.

Eyes small, placed laterally on head. Mouth small, terminal, oblique. Frenum absent.

Barbels absent. Pharyngeal tooth pattern 0,4-4,0. Gill rakers narrow and short. Dorsal fin with 8 rays and a slightly rounded distal end. Caudal peduncle short, thick. Caudal fin forked. Anal fin moderately large with 9 rays. Pelvic fins with 8 rays; insertion slightly anterior to dorsal fin insertion. Pectoral fins generally with 14 rays. Lateral line complete, slightly decurved with 32-36 cycloid scales in series. Spawning males develop a more intense iridescent blue shade on their dorsal and lateral sides as well as a defined, darker blue to purplish crescent mark posterior of the head. Spawning males also develop more intense orangish-red coloration on the dorsal side of their head as well as on the caudal, pelvic and pectoral fins. Tubercles are also present on the dorsal side of the head and on the snout of spawning males. Spawning females develop smaller tubercles on the head, and much less intense coloration.

Similar Species: Resembles the Common Shiner, Spotfin Shiner, and spawning Topeka

Shiner males. Common Shiner are generally larger in size, have a large eye, and large 184 diamond shaped scales that are nearly three times greater in depth than their width.

Spotfin Shiner have darker pigment forming a spot that is present on the second or third posterior rays of the dorsal fin. Spawning Topeka Shiner males can be distinguished from

Red Shiner by the presence of a dusky gray lateral stripe extending from the caudal peduncle to the snout and a lateral line that is outlined with small, paired, dark specks or

“mouse tracks”.

Distribution and Habitat: Native to North America in the Mississippi River basin from

Wyoming and South Dakota in the west, to southern Wisconsin and eastern Indiana in the east, and south throughout the Gulf drainages west of the Mississippi in Louisiana, Texas and New Mexico.13 The species is also widely introduced and distributed outside of its native range in the U.S.16 Present in the majority of the eastern and western main tributary basins of the Missouri River in South Dakota, as well as the Minnesota River basin. In North Dakota, the species has been documented throughout the Red River of the

North, James, Missouri, Cannonball, Knife and Heart River basins. Red Shiner are habitat generalists and can adapt to many habitat types. The species can be found in creeks, small to large streams, rivers, reservoirs, ponds and lakes. Red Shiner are rather tolerant of harsh environmental conditions including high turbidity, siltation, low or intermittent flows and increased temperatures.1,15 The species can often be found in pools with rocky substrates, or in runs and riffles of creeks and small rivers with sand, gravel and silt substrates.1,16 In the lower Platte River, Nebraska, Red Shiners selected areas with depths <30 cm (0.39 in) and velocities of <30 cm/s.21 Due to their aggressive behavior and habits, Red Shiner are capable of negatively affecting the populations, gene pools, and distributions of other native fish species.7,9,12 185

Reproduction: Spawning season is rather extended, and can take place between late

April and October when waters reach 15.6-29.4°C (60-85°F), which contributes to the species ability to rapidly increase in number.1,2,3,6,11 Spawning occurs intermittently and can take place within a variety of habitats such as over riffles and vegetation, within crevices, or over nests constructed by other fish such as sunfish.4,5,10,14,19 Sexual maturity is reached as early as the first summer of life.8 In Georgia, the smallest mature females collected from streams measured 30.5 mm (1.20 in) SL.5 Fecundity is not known to be dependent of the length and weight of females. Hatching occurs within 3-4 days.18

Age and Growth: Growth within the first season is known to be rapid.20 Larvae in a laboratory setting were reported attaining 16.4 mm (0.65 in) TL at 34 days post hatching.18 Mean lengths-at-age of males from the lower Platte River in Nebraska are reported as: age-0, 29.9 mm (1.18 in) FL; age-1, 35.6 mm (1.40 in) FL; age-2, 46.0 mm

(1.81 in) FL; age-3, 52.3 mm (2.06 in) FL; age-4, 57.0 mm (2.24 in) FL.20 Females in the

Platte River were longer and heavier than males in the earlier stages of life, but in the later years males showed to be longer and heavier than females.20 Capable of reaching 90 mm (3.54 in) TL.14 Longevity 4 years.20

Food and Feeding: Red Shiner have opportunistic, generalist, and omnivorous feeding characteristics. Feeding takes place by sight and is known to occurs throughout all depths in which it inhabits.1 Main food items include terrestrial and aquatic insects and larvae, plankton, and algae.1 Red Shiner have also been known to consume native larval fish in areas where they have been introduced.17

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 186

2. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

3. Farringer III, R.T., A.A. Echelle, and S.F. Lehtinen. 1979. Reproductive cycle of

the Red Shiner, Notropis lutrensis, in central Texas and south central Oklahoma.

Transactions of the American Fisheries Society 108:271-276.

4. Gale, W.F. 1986. Intermediate fecundity and spawning behavior of captive Red

Shiners-fractional, crevice spawners. Transactions of the American Fisheries

Society 115:429-437.

5. Herrington, S.J., and D.R. DeVries. 2008. Reproductive and early life history of

nonindigenous Red Shiner in the Chattahoochee River drainage, Georgia.

Southwestern Naturalist 7:413-428.

6. Jennings, M.R., and M.K. Saiki. 1990. Establishment of Red Shiner, Notropis

lutrensis, in the San Joaquin Valley, California. California Fish and Game 76:46-

57.

7. Karp, C.A., and H.M. Tyus. 1990. Behavioral interactions between young

Colorado squawfish and six fish species. Copeia 1990: 25-34.

8. Marsh-Matthews, E., W.J. Matthews, K.B. Gido, and R.L. Marsh. 2002.

Reproduction by young-of-year Red Shiner (Cyprinella lutrensis) and its

implications for invasion success. Southwestern Naturalist 47:605-610.

9. Mayden, R.L. 1989. Phylogenetic studies of North American minnows, with

emphasis on the genus Cyprinella (Teleostei: Cypriniformes). University of

Kansas Museum of Natural History, Miscellaneous Publication 80, Lawrence,

Kansas. 187

10. Minckley, W.L. 1972. Notes on the spawning behavior of Red Shiner, introduced

into Burro Creek, Arizona. Southwestern Naturalist 17:101-103.

11. Moyle, P.B. 2002. Inland fishes of California. University of California Press, Los

Angeles, CA. 612pp.

12. Nico, L., P. Fuller, and M. Neilson. 2014. Cyprinella lutrensis. USGS

Nonindigenous Aquatic Species Database, Gainesville, Florida. Available:

http://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=518.

13. Page, L.M. and B.M. Burr. 1991. A field guide to freshwater fishes of North

America north of Mexico. The Peterson Guide Series, vol. 42. Houghton Mifflin

Company, Boston, MA.

14. Pflieger, W.L. 1997. The Fishes of Missouri (revised). Missouri Department of

Conservation, Jefferson City, MO. 343 pp.

15. Poulos, H.M., B. Chernoff, P.L. Fuller, and D. Butman. 2012. Mapping the

potential distribution of the invasive Red Shiner, Cyprinella lutrensis (Teleostei:

Cyprinidae) across waterways of the conterminous United States. Aquatic

Invasions 7: 377-385.

16. Robins, C.R., R.M. Bailey, C.E. Bond, J.R. Brooker, E.A. Lachner, R.N. Lea, and

W.B. Scott. 1991. Common and scientific names of fishes from the United States

and Canada. American Fisheries Society Special Publication No. 20.

17. Ruppert, J.B., R.T. Muth, and T.P. Nesler. 1993. Predation on fish larvae by adult

red shiner, Yampa and Green Rivers, Colorado. The Southwestern Naturalist

38:397-399. 188

18. Saksena, V.P. 1962. The post-hatching stages of Red Shiner, Notropis lutrensis.

Copeia 1962:539-544.

19. Smith, P.W. 1979. The fishes of Illinois. University of Illinois Press, Urbana, IL.

314 pp.

20. Yildirim, A., and E.J. Peters. 2006. Life history characteristics of Red Shiner,

Cyprinella lutrensis, in the lower Platte River, Nebraska, USA. Journal of

Freshwater Ecology 21:307-314.

21. Yu, S., and E.J. Peters. 2002. Diel and seasonal habitat use by red shiner

(Cyprinella lutrensis). Zoological Studies 41:229-235.

189

Spotfin Shiner, Cyprinella spiloptera (Cope, 1867)

Etymology and Synonyms: Cyprinella = Latin for “cyprinus”, meaning “carp”; spiloptera = “spilos”, meaning “spot” and, “pteron”, meaning “wing” or “fin”, referring to the dark spot present of the dorsal fin.

Description: Body fusiform, moderately deep and laterally compressed; body depth goes more than 3.5 times into the SL. Dorsally dark to dusky gray; laterally silver, sometimes with a faint iridescent bluish-purple tint; ventrally silver to white; fins transparent with a single dark spot present on the second or third posterior rays of the dorsal fin. Head small. Snout short, tapering to a blunt point. Eyes small, placed laterally on head. Mouth large, terminal and strongly oblique; upper jaw extends below posterior nostril. Frenum absent. Barbels absent. Pharyngeal tooth pattern 1,4-4,1. Gill rakers short and conical; roughly 8-10. Dorsal fin with 8 rays. Caudal peduncle slightly elongate and uniform in thickness. Caudal fin forked. Anal fin moderately large with 8-9 rays. Pelvic fins abdominal with 8 rays; insertion slightly anterior to the dorsal fin insertion. Pectoral fins with 13-15 rays. Lateral line complete and slightly decurved with 35-39 cycloid scales in series. Nuptial males develop a more intense shade of blue on their dorsal and lateral sides, as well as a dusky stripe on their posterior lateral sides. Spawning males also develop yellowish-orange pigmentation on the anal, pelvic and pectoral fins, as well as tubercles on the snout, dorsal side of the head, lower jaw and pectoral fins. Spawning females develop occasional small tubercles on the dorsal side of the head and on the mid- dorsal scales.

Similar Species: Resembles the Red Shiner, Common Shiner and Sand Shiner. Red

Shiner have a deeper body with a body depth that goes less than 3.5 times into the 190 standard length. Red Shiner also lack a small dark spot of the dorsal fin. Common Shiner are generally larger in size, have a large eye, and large diamond shaped scales that are nearly three times greater in depth than their width. Sand Shiner have dorsal scales outlined in darker pigment creating a “crosshatched” pattern, and have a prominent, narrow mid-dorsal stripe present that widens into a wedge-shaped spot at the anterior base of the dorsal fin. Sand Shiner also have small, paired, dark specks or “mouse tracks” outlining the lateral line.

Distribution and Habitat: The native range of Spotfin Shiner consists of the Atlantic

Slope and St. Lawrence drainages from New York to eastern North Carolina in the east, throughout the Great Lakes, Hudson Bay and Mississippi River drainages in the west from eastern North Dakota south to eastern Oklahoma.7 Spotfin Shiner have been documented to occur within the Red River of the North and its major tributaries, as well as the Sheyenne, James, Cannonball, Heart, and Knife River drainages in North Dakota.

In South Dakota, Spotfin Shiner have a more limited distribution and are known to occur within the Big Sioux, Vermillion, James, and Missouri River drainages in the very southeastern part of the state. Spotfin Shiner inhabit shallower depths of medium to large sized river and streams in riffle or pooled areas, as well as larger lakes and reservoirs like

Louis and Clark Lake in South Dakota.1,6 The species is known to be rather tolerant of turbid waters with higher siltation levels, and is most frequently found over sand, mud, and gravel substrates.1

Reproduction: The spawning period of Spotfin Shiner takes place during the summer and is capable of extending from May to September.1,3 In Wisconsin, spawning activity was most frequent in water temperatures of 21.1-23.9°C (69.98-75.02°F).1 Spotfin Shiner 191 are fractional spawners, and also are known to spawn within crevices in rocky substrate.4

Male Spotfin Shiners are known to aggressively defend their smaller spawning territories from other spawning males.1,4,8 When two spawning males encounter one another within ones spawning territory, they can often be seen with erect dorsal fins swimming alongside one another, and sometimes nudging one another’s side with their snout, or grabbing the anal or pelvic fin with their mouth.1,4,8 When males encounter a female within their spawning territory, the are known to make several prespawning passes or movements before they go up and initiate spawning.1,4 Once females are lured by the male into the spawning territory or crevice, the pair will vibrate their bodies together to initiate the release of eggs and milt.4,8 It has been reported that as many as 10-97 eggs are released during a single spawning occurrence, with as many as 169-945 eggs deposited during a single spawning season.1,4 Eggs are adhesive, roughly 1.2-1.5 mm (0.05-0.06 in) in diameter, and hatch within roughly five days in water temperatures near 22°C

(71.6°F).1,4,5

Age and Growth: Information on Spotfin Shiner age and growth within the Dakotas region is scarce. In Wisconsin, mean lengths-at-age are reported as: age-1, 53.3 mm (2.09 in) TL; age-2, 62.1 mm (2.44 in) TL; age-3, 75.2 mm (2.96 in) TL.1 Males are known to be larger than females at each age, however females are known to live longer than males.1 Capable of reaching 108 mm (4.25 in) TL.9 Longevity 4 years.

Food and Feeding: Spotfin Shiner are opportunistic omnivores and feed diurnally on a variety of aquatic insects and larvae such as Trichoptera, and Diptera larvae, as well as terrestrial insects, small fish, algae or vegetative matter, and even their own eggs during spawning.1,2,4,8,10 192

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Cross, F.B., and J.T. Collins. Fishes in Kansas. 1995. Natural History Museum,

University of Kansas.

4. Gale, W.F., and C.A. Gale. 1977. Spawning habits of Spotfin Shiner (Notropis

spilopterus) – a fractional crevice spawner. Transactions of the American

Fisheries Society 100:170-177.

5. Hubbs, C.L., and G.P. Cooper. 1936. Minnows of Michigan. Cranbrook Institute

of Science Bulletin No. 8. 95pp.

6. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

7. Page, L.M., and B.M. Burr. 1991. A field guide to freshwater fishes: North

America north of Mexico. Houghton Mifflin Harcourt.

8. Pflieger, W.L. 1965. Reproductive behavior of the minnows Notropis spilopterus

and Notropis whipplii. Copeia 1965:1-8.

9. Stone, U.B. 1940. Studies on the biology of the satinfin minnows, Notropis

analostanus and Notropis spilopterus. Cornell University, Ithaca. PhD Thesis

98pp. 193

10. White, S.T., and D.C. Wallace. 1973. Diel changes in the feeding activity and

food habits of the spotfin shiner, Notropis spilopterus (Cope). The American

Midland Naturalist 90:200-205.

194

Common Carp, Cyprinus carpio (Linnaeus, 1758)

Etymology and Synonyms: Cyprinus = old Greek name for “carp”; carpio = Latinized name for “carp”.

Description: Body robust, arched back, deep, laterally compressed. Dorsally dark olive- gray to bronze; laterally golden yellow to light yellow-tan or olive with scales outlined in darker color appearing cross-hatched; ventrally light yellow to cream; pectoral, pelvic, anal, and caudal fins often yellow-orange in color on large adults. Head conical, moderately large. Snout long. Eye moderately large, placed laterally on upper portion of head. Mouth terminal to slightly subterminal; jaws do not extend backward to anterior edge of eye. Barbels present; 1 short barbel on each side of snout; 1 longer barbel on each corner of mouth. Lips thin. Teeth on jaws absent. Pharyngeal teeth molariform, aligned in

3 rows, 1,1,3-3,1,1. Gill rakers 21-27. Dorsal fin with 1 stiff, serrated spinous ray anteriorly, followed by 15-23 soft rays. Adipose fin absent. Caudal peduncle short, thick.

Caudal fin forked. Anal fin with 4-6 rays, including one stiff spinous ray anteriorly serrated posteriorly. Pelvic fins abdominal with 8-9 rays. Pectoral fins with 14-17 rays; do not extend to insertion of pelvic fins. Lateral line complete with 32-41 large cycloid scales in series. Spawning males slightly darker in color and develop tubercles on head and pectoral fins. Juveniles with a deeper body and larger eyes than adults.

Similar Species: May be confused with Grass Carp, Goldfish, and Buffalo (Ictiobus) species. Grass Carp lack any barbels and have a more elongate body that is less laterally compressed and more cylindrical than Common Carp. Goldfish have a similar body shape, but lack any barbels. Bigmouth and Smallmouth Buffalo lack barbels and any spines in the dorsal and anal fins. 195

Distribution and Habitat: One of the most widely distributed and highly invasive fish species in the world. Introduced to every continent except Antarctica.9 Native to Eastern

Europe and central Asia. First introduced to the United States as means of a food source in the 1870’s and to South Dakota eastern lakes around 1885-1886.1,3 The U.S. Bureau of

Fish and Fisheries continued stocking Common Carp into southeastern South Dakota lakes until roughly 1914.1 Widely distributed across the Dakotas. Habitat generalist.

Successful introductions and wide distribution mainly due to the species high adaptability to a wide variety of warm water habitats. Often inhabit shallow, highly conductive lakes, reservoirs, streams, ponds, and sloughs with clear to turbid water and abundant aquatic vegetation over soft substrate.5,11 Less frequent in cold, clear waters. Prefer littoral habitats during spring and summer, and congregate to deeper waters during late fall and winter months.7,11 In an eastern South Dakota glacial lake, fish were most active during dusk periods and least active during dawn periods.7 The species has serious detrimental effects on biotic and abiotic factors of aquatic ecosystems through their feeding and spawning habits, often causing decreased water quality and increased turbidity levels, making living conditions difficult for native fish communities and aquatic plant growth.8,16,18 Increased Common Carp density is capable of shifting phytoplankton communities from the important green algae, to often toxic cyanobacteria.20 Extremely tolerant of stressful environmental conditions including high water temperature, low oxygen levels, and turbidity, often times allowing them to achieve high abundance.

Reproduction: Adults known to concentrate in spawning grounds well before spawning occurs.15 Spawning is temperature dependent and takes place late spring through early summer within shallow shoreline areas with abundant macrophytes in marshes, wetlands, 196 and floodplains.7,11,15 Optimum spawning temperature 19-23 °C (66.2-73.4 °F), with spawning ceasing at temperatures >28 °C (82.4 °F).15 Spawning act consists of a single female often accompanied by several males, splashing near the surface.6 Prolific, fractional, broadcast spawners. Sexual maturity occurs at age 2-3.10 Females may produce over 2,000,000 eggs, with fecundity dependent on size of female.15 No nest is constructed and no parental care is given. Following spawning, adults broadly distribute themselves between shallow and deep waters.7,11,15 Eggs roughly 0.9-2.0 mm (0.04-0.08 in) in diameter, and adhesive to submerged macrophytes and soft substrate.2 Eggs hatch within

3-16 days depending on temperature.2 Hatching in eastern South Dakota glacial lakes occurred over a duration of up to 47 days from mid-May to early July.12

Age and Growth: Growth rapid during early years, and variable among populations, with length-at-age often related to population density.4,13 Age-0 individuals from South

Dakota glacial lakes grew approximately 1.00 mm/day (0.04 in/day), with increased growth rates influenced by warm water temperatures.12 Individuals who hatched earlier in the season exhibited faster growth rates and achieved greater total lengths during the first growing season than individuals that hatched later.12 Size structure known to decrease, and relative abundance known to increase with increasing depth.19 Average length at age-

1 in South Dakota waters has been reported as 132 mm (5.20 in) TL in Lake Francis

Case, 97 mm (3.82 in) TL in Lewis and Clark Reservoir, and 90 mm (3.54 in) TL in Lake

Oahe.13 Capable of exceeding 22.68 kg (50 lbs). Rarely exceed 20 years of age, but a longevity of 27 years has been reported in South Dakota.17

Food and Feeding: Adults opportunistic and omnivorous benthivores. Adult diets consist of detritus, and benthic macroinvertebrates such as chironomids, annelids, 197 odonates, and amphipods. Feeds by suctioning sediment into its mouth, where they sort out desired food organisms, and eject any unwanted material.14 Large individuals are capable of penetrating up to 12 cm (4.72 in) down into the substrate to find food.10 This behavior often results in the uprooting of submerged aquatic macrophytes, causing sediment resuspension and high levels of turbidity. Juveniles <100 mm (3.94 in) TL mainly consume large-bodied zooplankton before shifting to benthic prey items.16

Literature Cited:

1. Barnes, M.E. 2007. Fish hatcheries and stocking practices: past and present. Pages

267-294 in C. Berry, K. Higgins, D. Willis, and S. Chipps, editors. History of

fisheries and fishing in South Dakota. South Dakota Department of Game, Fish

and Parks, Pierre.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Blackwell, B.G. 2007. Warm-water fish species. Pages 213-238 in C. Berry, K.

Higgins, D. Willis, and S. Chipps, editors. History of fisheries and fishing in

South Dakota. South Dakota Department of Game, Fish and Parks, Pierre.

4. Carlander, K.D. 1969. Handbook of freshwater fishery biology, Vol. 1. Iowa State

University Press, Ames.

5. Egertson, C.J., and J.A. Downing. 2004. Relationship of fish catch and

composition to water quality in a suite of agriculturally eutrophic lakes. Canadian

Journal of Fisheries and Aquatic Sciences 61:1784-1796.

6. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville. 198

7. Hennen, M.J., and M.L. Brown. 2014. Movement and spatial distribution of

common carp in a South Dakota glacial lake system: implications for

management and removal. North American Journal of Fisheries Management

34:1270-1281.

8. Kloskowski, J. 2011. Impact of common carp Cyprinus carpio on aquatic

communities: direct trophic effects versus habitat deterioration. Fundamental and

Applied Limnology 178:245-255.

9. McCrimmon, H.R. 1968. Carp in Canada. Fisheries Research Board of Canada

Bulletin 165:1-93.

10. Panek, F.M. 1987. Biology and ecology of carp. Pages 1-15 in E.L. Cooper,

editor. Carp in North America. American Fisheries Society, Bethesda, Maryland.

11. Penne, C.R., and C.L. Pierce. 2008. Seasonal distribution, aggregation, habitat

selection of common carp in Clear Lake, Iowa. Transactions of the American

Fisheries Society 137:1050-1062.

12. Phelps, Q.E., B.D.S. Graeb, and D.W. Willis. 2008. First year growth and survival

of common carp in two glacial lakes. Fisheries Management and Ecology 15:85-

91.

13. Shields, J.T. 1970. Changes in the fish population in Lake Francis Case in South

Dakota in the first 16 years of impoundment. U.S. Bureau of Sport Fisheries and

Wildlife, Technical Paper 56, Washington, D.C.

14. Sibbing, F.A. 1988. Specialization and limitations in the utilization of food

resources by the carp, Cyprinus carpio: A study of oral food processing.

Environmental Biology of Fishes 22:161-178. 199

15. Swee, U.B., and H.R. McCrimmon. 1966. Reproductive biology of the carp,

Cyprinus carpio L., in Lake St. Lawrence, Ontario. Transactions of the American

Fisheries Society 95:372-380.

16. Weber, M.J., and M.L. Brown. 2009. Effects of common carp on aquatic

ecosystems 80 years after “carp as a dominant”: ecological insights for fisheries

management. Reviews in Fisheries Science 17:524-537.

17. Weber, M.J., and M.L. Brown. 2011a. Comparison of common carp (Cyprinus

carpio) age estimates derived from dorsal fin spines and pectoral fin rays. Journal

of Freshwater Ecology 26:195-202.

18. Weber, M.J., and M.L. Brown. 2011b. Relationships among invasive common

carp, native fishes and physiochemical characteristics in upper Midwest (USA)

lakes. Ecology of Freshwater Fish 20:270-278.

19. Weber, M.J., M.L. Brown, and D.W. Willis. 2010. Spatial variability of common

carp populations in relation to lake morphology and physiochemical parameters in

the upper Midwest United States. Ecology of Freshwater Fish 19:555-565.

20. Williams, A.E., and B. Moss. 2003. Effects of different fish species and biomass

on plankton interactions in a shallow lake. Hydrobiologia, 491:331-346.

200

Western Silvery Minnow, Hybognathus argyritis (Girard, 1856)

Etymology and Synonyms: Hybognathus = “swollen jaw”; argyritis = unknown.

Description: Body fusiform, slightly elongate, moderately laterally compressed. Dorsally tan-olive to silver; laterally silver with no obvious markings; ventrally silver to white; fins clear, transparent. Head broad; length less than 5x the diameter of the eye.

Basioccipital process internally posterior of head is broad, straight and barely concave; muscles well separated at point of attachment. Snout elongate, slightly protrudes past mouth. Eye moderately large, placed laterally on head. Mouth subterminal, small, crescent shaped. Frenum absent. Barbels absent. Lips thin. Pharyngeal tooth pattern 0,4-

4,0. Dorsal fin with 8 rays, straight distal end and pointed tip. Adipose fin absent. Caudal peduncle slightly elongate, moderately thick. Caudal fin forked. Anal fin with 8 rays.

Pelvic fins abdominal; insertions slightly posterior to dorsal fin insertion. Lateral line complete with 36-40 large cycloid scales in series. Spawning males develop small tubercles of dorsal side of head, opercles, and pectoral fins.

Similar Species: Closely resembles the Plains Minnow. Plains Minnow have a smaller eye, and a basioccipital process that is narrow and peg-like with muscles almost touching the point of attachment. Brassy Minnow have a dorsal fin with a distinctly rounded tip, and have small black specks present on the caudal fin. Flathead Chub have a broad, dorsoventrally flattened head tapering to a pointed snout, barbels, and have sickle shaped dorsal and pectoral fins.

Distribution and Habitat: Native to the Missouri and Red River of the North river drainages. Primarily found within the main tributaries west of the Missouri River in the

Dakotas. Once widespread throughout the Great Plains region, the abundance and 201 distribution of the species has been in decline across their historic range, in part due to irrigation and impoundments, which reduce turbidity and increase the proportion of large substrates.2,4,9,10,13,14 Western Silvery Minnow are adapted to the hydrologically dynamic and turbid conditions, common in prairie streams and rivers throughout the Great Plains, and occurs most frequently in channel border habitat and areas with little to no current such as backwaters and pools over silt and sand substrates.3,7,10,15,16 In the Missouri and

Yellowstone Rivers in North Dakota, the species was most frequently taken from depths less than 1 m (3.28 ft) with current velocities less than 0.5 m/s (1.6 ft/s).16 Most individuals were also captured in areas with turbidities less than 250 NTU and in water temperatures 18-22 °C (64.4-71.6 °F).16 Avoids reaches with gravel and large rocky substrate, as well as areas with an abundance of piscivores.10 Fry and juveniles are often found schooling within quiet inshore areas and at the mouths of tributaries nearly two weeks after hatching.9,11

Reproduction: Information regarding the spawning and reproductive habits of Western

Silvery Minnow is extremely scarce or unknown. Individuals from Missouri are known to be in spawning condition during late June, and in Alberta are presumed to spawn in

May.8,12 An early study from New York on the closely related Eastern Silvery Minnow is the only available information regarding fecundity, spawning and early development.9,11

Spawning migrations consist of movements made to inshore waters of lakes and larger rivers in the spring.11 Females likely reach sexual maturity at age-1, or at 50-55 mm

(1.97-2.17 in) SL.11 Males mature later, likely at age-2.11 Males are not known to defend territories or show aggressiveness towards other mature males.11 Spawning takes place during the day in shallow waters with abundant aquatic vegetation and ceases at 202 night.5,9,11 One to ten males initially pursue a single female until two males establish themselves on each side of the female and swim alongside her.11 The males have been noted to nudge the female in the abdominal region with their snouts, and rapidly vibrate their bodies together to initiate the release of eggs and milt.11 Females retreat to deeper waters post spawning while males continue to look for other mature females.11 No nest is prepared and no parental care is given. Fecundity increases with size of female.11 Eggs roughly 1 mm (0.04 in) in diameter, milky in color, non-adhesive and demersal.11

Hatching takes place within 6-7 days.11

Age and Growth: Larvae average 5.5 mm (0.22 in) SL and 6.0 mm (0.24 in) TL at hatching.11 Mean lengths-at-age from the Moreau River in South Dakota are reported as: age-1, 70 mm (2.76 in) TL; age-2, 109 mm (4.29 in) TL; age-3, 119 mm (4.69 in) TL; age-4, 151 mm (5.95 in) TL.6 Capable of reaching 152.4 mm (6 in) TL.1 Longevity 5.5 years.8

Food and Feeding: Information on the diet of the Western Silvery Minnow is vastly unknown, but is presumed to be similar to the Plains Minnow, Eastern Silvery Minnow and Central Silvery Minnow.1,7,9,11 A long, coiled intestine and a crescent shaped subterminal mouth allow the species to consume large amounts of microscopic vegetative matter, diatoms, organic materials and fungi.11,17

Literature Cited:

1. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

2. Cross, F.B., R.L. Mayden, and J.D. Stewart. 1986. Fishes in the western

Mississippi drainage. Pages 363-412 in C.H. Hocutt and E.O. Wiley (editors). The 203

Zoogeography of North American Freshwater Fishes. John Wiley and Sons,

Toronto, Canada.

3. Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, 5.

Selected chubs and minnows (Cyprinidae): sicklefin chub (M. meeki), sturgeon

chub (M. gelida), silver chub (M. storeriana), speckled chub (M. aestivaus),

flathead chub (Platygobio gracilis), plains minnow (Hybognathus placitus), and

western silvery minnow (H. argyritis). Transactions of the Nebraska Academy of

Sciences and Affiliated Societies 21:99-108.

4. Hesse, L.W., J.C. Schmulbach, J.M. Carr, K.D. Keenlyne, D.G. Unkenholz, J.W.

Robinson, and G.E. Mestl. 1989. Missouri River fishery resources in relation to

past, present, and future stresses. Pages 352-371 in D.P. Dodge, editor.

Proceedings of the international large river symposium. Canadian Special

Publication of Fisheries and Aquatic Sciences 106.

5. Houston, J. 1998. Status of the western silvery minnow, Hybognathus argyritis, in

Canada. Canadian Field-Naturalist 112:174-153.

6. Jones, S.J. 2018. Western prairie stream fisheries: an assessment of past and

present fish assemblage structure, biotic homogenization, and population

dynamics in western South Dakota streams. M.S. Thesis, South Dakota State

University, Brookings, South Dakota.

7. Pflieger, W.L. 1980. Hybognathus argyritis Girard, western silvery minnow. Page

174 in D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and

J.R. Stauffer Jr. (editors). Atlas of North American Freshwater Fishes. North 204

Carolina State Museum of Natural History, North Carolina Biological Survey

Publication 1980-12.

8. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of

Conservation, Jefferson City, Missouri.

9. Pollard, S.M. 2003. Status of the western silvery minnow (Hybognathus argyritis)

in Alberta. Alberta Wildlife Status Report No. 47.

10. Quist, M.C, W.A. Hubert, and F.J. Rahel. 2004. Relations among habitat

characteristics, exotic species, and turbid-river cyprinids in the Missouri River

drainage of Wyoming. Transactions of the American Fisheries Society 133:727-

742.

11. Raney, E.C. 1939. The breeding habits of the eastern silvery minnow,

Hybognathus regius Girard. American Midland Naturalist 21:215-218.

12. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

13. Smith, C.D., J.R. Fischer, and M.C. Quist. 2014. Historical changes in Nebraska’s

lotic fish assemblages: implications of anthropogenic alterations. The American

Midland Naturalist 172:160-184.

14. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67. 205

15. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press, Columbus, Ohio.

16. Welker, T.L., and D.L. Scarnecchia. 2004. Habitat use and population structure of

four native minnows (family Cyprinidae) in the upper Missouri and lower

Yellowstone rivers, North Dakota (USA). Ecology of Freshwater Fish 13:8-22.

17. Whitaker, J.O. 1977. Seasonal changes in food habits of some cyprinid fishes

from the White River at Petersburg, Indiana. The American Midland Naturalist

90:411-418.

206

Brassy Minnow, Hybognathus hankinsoni (Hubbs, 1929)

Etymology and Synonyms: Hybognathus = “swollen jaw”; hankinsoni = named after naturalist and ichthyologist T.L. Hankinson of Michigan.

Description: Body fusiform, robust, slightly laterally compressed. Dorsally dark olive to bronze with faint, dark mid-dorsal stripe; laterally olive to bronze or silver with thick, dark lateral stripe that may appear iridescent purple; ventrally cream to white; fins light olive to bronze in color; rays of lower lobe of caudal fin with small black specks present.

Head short. Snout bluntly rounded, short, slightly protruding past mouth. Basioccipital process internally posterior of head is narrow, peg-like, and shorter than basioccipital process in Plains Minnow. Eye large, placed laterally on head. Mouth small, terminal, slightly oblique; upper jaw does not extend to anterior edge of eye. Barbels absent.

Frenum absent. Lips thin. Pharyngeal tooth pattern 0,4-4,0. Dorsal fin with 8 rays; anterior ray slightly shorter than the second or third rays; distal end slightly rounded.

Adipose fin absent. Caudal peduncle slightly elongate, uniform in thickness. Caudal fin moderately forked. Anal fin with 7-8 rays and straight distal end. Pelvic fins with 8 rays and rounded distal end; insertions slightly posterior to dorsal fin insertion. Pectoral fins with 13-15 rays. Lateral line complete with 35-39 cycloid scales in series. Spawning males gold or brassy in color with small tubercles present on the pectoral fins. Juveniles similar in appearance to adults.

Similar Species: Closely resembles the Plains Minnow and Western Silvery Minnow.

Plains Minnow are silver in color, have a smaller eye, a subterminal mouth, a shorter snout, and a dorsal fin with a straight distal end and pointed tip. Western Silvery Minnow 207 are silver in color, have a subterminal mouth, a shorter snout, and a dorsal fin with a straight distal end and pointed tip.

Distribution and Habitat: Native to southern Canada from British Columbia to Quebec, and the north central United States from Montana to Colorado in the west, east throughout the Missouri, upper Mississippi, upper St. Lawrence and Great Lakes drainages to New York, and south to northern Missouri and Kansas. Primarily occurs throughout the main western tributaries of the Missouri River in North Dakota.

Widespread throughout the Missouri River basin in South Dakota, but is known to be absent from the Belle Fourche and Moreau watersheds. The native range has been declining as well as the abundance of the species, likely due to habitat fragmentation and altered hydrology, which can negatively affect annual recruitment.4,5,8 Often found schooling in creeks, headwater and narrow streams, small rivers, and lakes with larger substrates covered in silt or organic debris and underbank coverage in low velocity areas such as slow runs and pools.1,6,10,12 Generally avoids large rivers and lakes.2 Rapid dispersal is likely during wet seasons.10 During drought and overwintering periods, individuals often seek out deeper pools.5,10 Good predictor variables of Brassy Minnow presence is higher conductivity levels (101-2060 μS/cm), and highly productive waters with abundant submerged vegetation and high nutrient imputs.8 Capable of persisting in low oxygenated waters (as low as 0.03 mg/L) and in water temperatures as high as 35.5

°C (95.9 °F).10 When conditions of backwater areas deteriorate, larvae migrate to mid- channel habitats.5

Reproduction: Information on spawning and reproductive habits of Brassy Minnow is scarce. Spawning takes place during spring and early summer when water temperatures 208 reach 16-27 °C (60.8-80.6 °F), within quiet, seasonally flooded, shallow, vegetated backwater and floodplain habitats.3,5,10,11 Spawning and recruitment are strongly affected by intermittency and dewatering events such as stream geomorphology, groundwater pumping, and climate.5 Sexual maturity occurs between ages 1-2.2,3 Spawning activity has been recorded to take place in the afternoon, and consist of one or more males pursuing a single female, and pressing against her with the pair vibrating their bodies to initiate the release of eggs and milt.3 Mature eggs likely yellow in color and roughly 0.7 mm (0.03 in) in diameter.2 Eggs are adhesive to vegetation and other submerged structure.5 Hatching occurs within 3 days.5

Age and Growth: Larvae approximately 4.0 mm (0.16 in) SL at hatching.5 Lengths of individuals from southeastern Wyoming are reported as: age-0, 24-44 mm (0.94-1.73 in)

TL; age-1, 44-68 mm (1.73-2.68 in) TL; age-2, 66-80 mm (2.60-3.15 in) TL; age-3, 82-

84 mm (3.23-3.31 in) TL.3 Growth is not known to be significantly different between sexes.3 Growth rates have been shown to be strongly affected by habitat drying.5 Little growth occurs during winter months.3 Capable of reaching 97 mm (3.82 in) TL.7

Longevity 4 years.9

Food and Feeding: Opportunistic omnivore.3 Often found foraging in large schools slowly moving upstream, with foraging increasing during periods of high water.3 Feeds heavily on phytoplankton, diatoms, algae and organic debris.3 Also consumes small aquatic invertebrates including diptera larvae and microcrustaceans such as copepods and cladocerans.3

Literature Cited: 209

1. Bailey, R.M. 1954. Distribution of the American cyprinid fish Hybognathus

hankinsoni with comments on its original description. Copeia 1954:289-291.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Copes, F.A. 1975. Ecology of the brassy minnow Hybognathus hankinsoni

(Cyprinidae). Reports on the Flora and Fauna of Wisconsin 10:46-72.

4. Falke, J.A., K.D. Fausch, R. Magelky, A. Aldred, D.S. Durnford, L.K. Riley, and

R. Oad. 2011. The role of groundwater pumping and drought in shaping

ecological futures for stream fishes in a dryland river basin of the western Great

Plains, USA. Ecohydrology 4:682-697.

5. Falke, J.A., K.R. Bestgen, and K.D. Fausch. 2010. Streamflow reductions and

habitat drying affect growth, survival, and recruitment of brassy minnow across a

Great Plains riverscape. Transactions of the American Fisheries Society

139:1566-1583.

6. Fischer, J.R., and C.P. Paukert. 2008. Habitat relationships with fish assemblages

in minimally disturbed Great Plains regions. Ecology of Freshwater Fish 17:597-

609.

7. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

8. Nowosad, D.M., and E.B. Taylor. 2013. Habitat variation and invasive species as

factors influencing the distribution of native fishes in the lower Fraser River

Valley, British Columbia, with an emphasis on brassy minnow (Hybognathus

hankinsoni). Canadian Journal of Zoology 91:71-81. 210

9. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

10. Scheurer, J.A., and K.D. Fausch. 2003. Multiscale processes regulate brassy

minnow persistence in a Great Plains river. Transactions of the American

Fisheries Society 132:840-855.

11. Starrett, W.C. 1951. Some factors affecting the abundance of minnows in the Des

Moines River, Iowa. Ecology 32:13-27.

12. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67.

211

Plains Minnow, Hybognathus placitus (Girard, 1856)

Etymology and Synonyms: Hybognathus = “swollen jaw”; placitus = “broad surface”; possibly referring to the broad head of the species.

Description: Body fusiform, slightly elongate, moderately laterally compressed. Dorsally yellowish-tan to silver with dusky dark olive mid-dorsal stripe; laterally silver with no obvious markings; ventrally silver to white; fins clear, transparent. Head broad, slightly ventrally flattened; length greater than or equal to 5x the diameter of the eye.

Basioccipital process internally posterior of head narrow and peg-like; muscles almost touch at point of attachment. Snout elongate, slightly protrudes past mouth. Eye moderately small, placed laterally on head. Mouth subterminal, small, crescent shaped.

Frenum absent. Barbels absent. Lips thin. Pharyngeal tooth pattern 0,4-4,0. Dorsal fin with 8 rays, straight distal end and pointed tip. Adipose fin absent. Caudal peduncle slightly elongate, moderately thick. Caudal fin forked. Anal fin with 8 rays. Pelvic fins abdominal; insertions slightly posterior of dorsal fin insertion. Lateral line complete with

36-40 large cycloid scales in series. Sexually dimorphic species; males display a longer, deeper head than females, and have an anterior dorsal fin ray that is longer than the rest of the dorsal fin rays when depressed; females generally have deeper bodies than males and have an anterior dorsal fin ray that is equal to the rest of the dorsal fin rays when depressed. Spawning males develop small tubercles on dorsal side of head, opercles, and pectoral fins.

Similar Species: Closely resembles the Western Silvery Minnow. Western Silvery

Minnow have a larger eye, and a basioccipital process that is broad, straight, and barely concave with muscles well separated at the point of attachment. Brassy Minnow have a 212 dorsal fin with a distinctly rounded tip and have small black specks present on the caudal fin. Flathead Chub have a broad, dorsoventrally flattened head tapering to a pointed snout, barbels, and have sickle shaped dorsal and pectoral fins.

Distribution and Habitat: Native range in the United States extends from Montana and

North Dakota in the north, south through the Great Plains to Texas.15 Historically considered to be one of the most abundant fishes in the Missouri River, the species abundance is in recent decline.4,5,14 Primarily found within the main tributaries west of the Missouri River in the Dakotas. Inhabits medium to large, often turbid rivers and streams with sand or silt substrates, some current, and exposed, shallow, sand-filled channels.3,11,15 Often found colonizing within shallow, slow water or pool habitats where organic debris accumulates.1,4 In a laboratory setting, individuals held at dissolved oxygen levels of 5 mg/L and above selected water temperatures near 30 °C (86 °F).2 In water with a dissolved oxygen level of 2 mg/L, the selected water temperature was only

17 °C (62.6 °F).2 This suggests that in low oxygenated waters, the species selects higher water temperatures, but less than 30 °C (86 °F), at which it can operate within the zone of respiratory independence.2 Mean critical thermal maximum 39.95 °C (103.91 °F).8

Mortality occurs in waters with salinities <22 ppt.10

Reproduction: Prolonged spawning season extends from late April to August, and is known to coincide with high or receding flows.3,7,15,16 Sexual maturity reached at ages 1-

2, or when individuals reach roughly 45-50 mm (1.77-1.97 in) SL.7,15 If preferred spawning conditions are infrequent, or do not occur in late summer, smaller mature individuals may skip spawning at age-1, and wait to spawn until the following spring.15

Fractional, pelagic, and broadcast spawner.7,12,16 Spawning takes place within quiet 213 backwaters along sandbars during receding flows.15 Males pursuing a single female nudge her abdominal region, and eventually will wrap their bodies around the female to initiate the release of eggs and milt.12 No nest is constructed and no parental care is given.

High levels of post-spawning mortality are common.11 Fecundity increases with size of female.15 In Oklahoma, fecundity ranged 417-4134 eggs from females 51-87 mm (2.01-

3.43 in) SL.15 Eggs roughly 1 mm (0.04 in) in diameter, semibuoyant, slightly demersal and non-adhesive allowing them to develop and drift down stream until hatching.9,12,13

Reproductive success is likely negatively impacted by stream fragmentation, since eggs and larvae require a certain unimpeded distance of river to develop.16 Given this requirement, periods of increased discharge are needed during the spawning season for successful spawning events.16

Age and Growth: Mean lengths-at-age from the Moreau River in South Dakota are reported as: age-1, 62 mm (2.44 in) TL; age-2, 95 mm (3.74 in) TL; age-3, 104 mm (4.09 in) TL.5 Capable of reaching 125 mm (4.92 in) TL.6 Longevity 2 years.15

Food and Feeding: Herbivore. A long, coiled intestine and a crescent shaped subterminal mouth allow the species to consume large amounts of vegetative matter and diatoms.3 Feeds by scraping and scooping up algae off submerged macrophytes and bottom substrates.3,4

Literature Cited:

1. Baxter, G.T., and M.D. Stone. 1995. Fishes of Wyoming. Wyoming Game and

Fish Department, Cheyenne, Wyoming. 214

2. Bryan, J.D., L.G. Hill, and W.H. Neill. 1984. Interdependence of acute

temperature preference and respiration in the plains minnow. Transactions of the

American Fisheries Society 113:557-562.

3. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

4. Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, 5.

Selected chubs and minnows (Cyprinidae): sicklefin chub (M. meeki), sturgeon

chub (M. gelida), silver chub (M. storeriana), speckled chub (M. aestivaus),

flathead chub (Platygobio gracilis), plains minnow (Hybognathus placitus), and

western silvery minnow (H. argyritis). Transactions of the Nebraska Academy of

Sciences and Affiliated Societies 21:99-108.

5. Jones, S.J. 2018. Western prairie stream fisheries: an assessment of past and

present fish assemblage structure, biotic homogenization, and population

dynamics in western South Dakota streams. Unpublished M.S. Thesis, South

Dakota State University, Brookings, South Dakota.

6. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

State Museum of Natural History, Raleigh, North Carolina.

7. Lehtinen, S.F., and J.B. Layzer. 1988. Reproductive cycle of the plains minnow,

Hybognathus placitus (Cyprinidae), in the Cimarron river, Oklahoma. The

Southwestern Naturalist 33:27-33. 215

8. Matthews, W.J., and J.D. Maness. 1979. Critical thermal maxima, oxygen

tolerances and success of cyprinid fishes in a southwestern river. The American

Midland Naturalist 102:374-377.

9. Miller, R.J., and H.W. Robison. 1973. The fishes of Oklahoma. Oklahoma State

University Press, Stillwater, Oklahoma.

10. Ostrand, K.G., and G.R. Wilde. 2001. Temperature, dissolved oxygen, and

salinity tolerances of five prairie stream fishes and their role in explaining fish

assemblage patterns. Transactions of the American Fisheries Society 130:742-

749.

11. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of

Conservation, Jefferson City, Missouri.

12. Platania, S.P., and C.S. Altenbach. 1998. Reproductive strategies and egg types of

seven Rio Grande basin cyprinids. Copeia 1998:559-569.

13. Sliger, A.S. 1967. The embryology, egg structure, micropyle, and egg membranes

of the plains minnow Hybognathus placitus (Girard). M.S. Thesis, Oklahoma

State University, Stillwater, Oklahoma.

14. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67. 216

15. Taylor, C.M., and R.J. Miller. 1990. Reproductive ecology and population

structure of the plains minnow, Hybognathus placitus (Pisces: Cyprinidae), in

central Oklahoma. The American Midland Naturalist 123:32-39.

16. Urbanczyk, A.C. 2012. Reproductive ecology of the plains minnow Hybognathus

placitus in the Brazos River, Texas. M.S. Thesis, Texas Tech University,

Lubbock, Texas.

217

Silver Carp, Hypophthalmichthys molitrix (Valenciennes, 1844)

Etymology and Synonyms: Hypo = “under” or “below”, phthalm = “eye”, ichthys =

“fish”, in reference to the positioning of the eye; molitrix = “miller”, or “grinder”, possibly referring to the grinding ability of the pharyngeal apparatus.

Description: Body fusiform, deep, robust, moderately laterally compressed. Dorsally light gray to olive; laterally solid silver; ventrally silver to white; dorsal part of head gray to olive; fins light to dark gray, pelvic fins somewhat transparent to white. Head large, scaleless. Snout short. Eye placed laterally on lower anterior end of head. Mouth large, terminal; lower jaw protrudes forward and beyond upper jaw; upper jaw does not extend to anterior end of eye. Barbles absent. Lips fleshy. Teeth absent on jaws. Pharyngeal teeth

0,4-4,0. Gill rakers highly modified, fused and sponge-like. Dorsal fin with 1 smooth spinous ray and 8 soft rays. Adipose fin absent. Caudal peduncle thick, slightly elongate.

Caudal fin forked. Anal fin falcate with 1 smooth spinous ray and 12-14 soft rays. Pelvic fins abdominal with 7-8 rays; insertion anterior to insertion of dorsal fin. Keel extending from gills to vent. Pectoral fins slightly elongate, extending to insertion of pelvic fins; 1 posterior serrated spinous ray and 15-18 soft rays. Lateral line decurved and complete with 85-100 small cycloid scales in series. Spawning adults similar to non-spawners.

Juveniles similar to adults, but lack spinous rays in dorsal, anal, and pectoral fins.

Similar Species: Closely resembles the Bighead Carp. Bighead Carp are dark gray dorsally and have small, dark, irregular blotches along the lateral sides. Bighead Carp also display a ventral keel that extends from the pelvic fins to the vent, and elongate pectoral fins that extend past the insertion of the pelvic fins. Gizzard Shad may be confused with smaller Silver Carp, however Gizzard Shad lack a lateral line, have a long 218 filament on the dorsal fin, and have a dark to black spot on the upper portion of the gill cover. Grass and Common Carp have much larger cycloid scales.

Distribution and Habitat: Native to the major Pacific drainages of eastern Russia and south to northern Vietnam, but has been extensively introduced worldwide as means of food.2,7 First introduced to the United States by commercial fish producers to aquaculture facilities in Arkansas and Arizona in the early 1970’s where the species subsequently escaped and began to migrate up the tributaries of the Mississippi River.6,7 In the United

States, the species is now established in the Mississippi River and its major tributaries.7

Found within the James, Vermillion, and Big Sioux watersheds of the Dakotas.

Dispersion into these Missouri River watersheds has decreased or halted due to artificial and natural barriers.4 Primarily a large river species, often found in areas >3 m (9.84 ft) in depth with low velocity such as behind wing dikes, floodplains, or backwaters, but also inhabit the open waters of lakes, reservoirs, and ponds.7,19 Optimum temperature for larvae 26-33.5 °C (78.8-92.3 °F).7,13,14 Upper lethal temperature for larvae 43.5-46.5 °C

(110.3-115.7 °F).7,12 Rather tolerant of low temperatures.7 Overwinter in pool habitat with activity slowing at temperatures <4 °C (39.2 °F), and little movement occurring <2

°C (35.6 °F).7

Reproduction: Spawning takes place spring and early summer or when water temperatures reach 17-26.5 °C (62.6-79.7 °F).16 Age of maturation heavily depends on the growth rate during the first year of life.5 Sexual maturity occurs by age-3 in the species native range, however age-2 individuals have been reported from the middle

Mississippi River.7,19 High growth rates during the early stages of life in the middle

Mississippi River may allow females to mature earlier than males.19 Prolific spawners. 219

Spawning act takes place near the water’s surface and consists of splashing and chasing while eggs and sperm are extracted.7,8 Fecundity increases with size and age of female.

Fecundity of six age-2 females from the middle Mississippi River was 57,283-328,538 eggs, with an average of 156,312 eggs per female.19 No nest is constructed, and no parental care is given. Eggs semibuoyant and carried with the current as far as 500 km

(310.69 mi) downstream until hatching.3,7,9 Eggs roughly 1.4 mm (0.06 in) in diameter before fertilization, and 4.9-5.6 mm (0.19-0.22 in) in diameter after water hardening.1,7

Hatching occurs within 21-33 hours in warmer waters.1

Age and Growth: Maximum growth occurs in water 24-34 °C (75.2-93.2 °F).7,10 Larvae roughly 4.60-5.55 mm (0.18-0.22 in) in length at hatching.1 Capable of reaching 400 mm

(15.75 in) TL and 270 g (0.60 lbs) by the end of their first growing season, and can gain

45 g (0.10 lbs) or more per month.4,17,18 Fast initial growth rates followed by slower growth rates with increasing age.4,19 Growth primarily influenced by food availability.7

Mean lengths-at-age from the middle Mississippi River are reported as: age-1, 317.7 mm

(12.51 in) TL; age-2, 530.9 mm (20.90 in) TL; age-3, 649.8 mm (25.58 in) TL; age-4,

704.1 mm (27.72 in) TL; age-5, 723.3 mm (28.48 in) TL.19 Females generally heavier than males.19 Capable of reaching 1260 mm (49.61 in) TL.7 Longevity 15+ years in its native range, but often reach age 5 in the Mississippi River and its tributaries.4,7,19

Food and Feeding: Opportunistic, planktivorous filter feeder. Larvae feed on small zooplankton such as rotifers and protozoans, with size of zooplankton increasing with the size of larvae.7,11 As individuals grow, diets shift from mainly zooplankton to phytoplankton.7 Adults primarily consume phytoplankton and zooplankton, and to a lesser extent detritus and bacteria.7 Silver Carp feeding habits often shift the species 220 composition of the phytoplankton and zooplankton communities from larger to smaller species, which have detrimental effects to other native fish species.7,15 Able to effectively consume smaller particles than Bighead Carp due to a smaller epibranchial organ.7 Active feeding takes place in water temperatures 15-30 °C (59-86 °F), but may continue to feed in waters as low as 4 °C (39.2 °F).7

Literature Cited:

1. Chapman, D.C., and A.E. George. 2011. Developmental rate and behavior of

early life stages of bighead and silver carp. U.S. Geological Survey Scientific

Investigations Report 2011-5076, 62p.

2. Fuller, P.L., L.G. Nico, and J.D. Williams. 1999. Non-indigenous fishes

introduced into inland waters of the United States. American Fisheries Society,

Special Publication 27, Bethesda, Maryland.

3. Gorbach, E.I., and M.L. Krykhtin. 1989. Migration of the white amur,

Ctenopharyngodon idella, and silver carp, Hypophthalmichthys molitrix, in the

Amur River Basin. Journal of Ichthyology 28:47-53.

4. Hayer, C.A., B.D.S. Graeb, and K.N. Bertrand. 2014. Adult, juvenile and young-

of-year bighead, Hypophthalmichthys nobilis (Richardson, 1845) and silver carp,

H. molitrix (Valenciennes, 1844) range expansion on the northwestern front of the

invasion in North America. BioInvasions Records 3:283-289.

5. Kamilov, B.G. 1987. Gonad condition of female silver carp Hypophthalmichthys

molitrix in relation to growth rate in Uzbekistan. Journal of Ichthyology 27:135-

139. 221

6. Kelley, A.M., C.R. Engle, M.L. Armstrong, M. Freeze, and A.J. Mitchell. 2011.

History of introduction and governmental involvement in promoting the use of

grass, silver, and bighead carps. Pages 163-174 in D.C. Chapman and M.H. Hoff,

editors. Invasive Asian carps in North America. American Fisheries Society,

Symposium 74, Bethesda, Maryland.

7. Kolar, C.S., D.C. Chapman, W.R. Courtenay, Jr., C.M. Housel, J.D. Williams,

and D.P. Jennings. 2007. Bighead carps: a biological synopsis and environmental

risk assessment. American Fisheries Society, Special Publication 33, Bethesda,

Maryland.

8. Kuronuma, K. 1968. New systems and new fishes for culture in the Far East. FAO

Fisheries Report 5:123.

9. Laird, C.A., and L.M. Page. 1996. Non-native fishes inhabiting the streams and

lakes of Illinois. Illinois Natural History Survey Bulletin 35:1-51.

10. Mahboob, S., and A.N. Sheri. 1997. Growth performance of major, common and

some Chinese carps under composite culture system with special reference to

pond fertilization. Journal of Aquaculture in the tropics 12:201-207.

11. Marciak, Z., and E. Bogdan. 1979. Food requirements of juvenile stages of grass

carp, Ctenopharyngodon idella Val., silver carp, Hypophthalmichthys molitrix

Val., and bullhead carp Aristichthys nobilis Rich. EMS Special Publication 4:139-

157.

12. Opuszynski, K., A. Lirski, L. Myszkowski, and J. Wolnicki. 1989. Upper lethal

and rearing temperatures for juvenile common carp, Cyprinus carpio L. and silver 222

carp, Hypophthalmichthys molitrix (Valenciennes). Aquaculture and Fisheries

Management 20:287-294.

13. Panov, D.A., and L.V. Khromov. 1970. Why the larvae die, Rybovodstvo I

Rypolovstvo 6:12-13. (In Polish).

14. Radenko, V.N., and I.A. Alimov. 1992. Significance of temperature and light for

growth and survival of larvae of silver carp, Hypophthalmichthys molitrix.

Journal of Ichthyology 32:16-27.

15. Sampson, S.J., J.H. Chick, and M.A. Pegg. 2009. Diet overlap among two Asian

carp and three native fishes in backwater lakes on the Illinois and Mississippi

rivers. Biological Invasions 11:483-496.

16. Schofield, P.J., J.D. Williams, L.G. Nico, P. Fuller, and M.R. Thomas. 2005.

Foreign nonindigenous carps and minnows (Cyprinidae) in the United Sates-a

guide to their identification, distribution and biology. US Geological Survey

Scientific Investigations Report 2005-5041.

17. Stone, N., C. Engle, D. Heikes, and D. Freeman. 2000. Bighead carp. Southern

Regional Aquaculture Center (SRAC), Stoneville, Mississippi, September 2000.

Southern Regional Aquaculture Center Publication 428.

18. Waterman, M.P. 1997. Chinese bighead carp continues to draw interest.

Aquaculture Magazine 23:73-79.

19. Williamson, C.J., and J.E. Garvey. 2005. Growth, fecundity, and diets of newly

established silver carp in the middle Mississippi River. Transactions of the

American Fisheries Society 134:1423-1430.

223

Bighead Carp, Hypophthalmichthys nobilis (Richardson, 1845)

Etymology and Synonyms: Hypo = “under” or “below”, phthalm = “eye”, ichthys =

“fish”, in reference to the positioning of the eye; nobilis = noble.

Description: Body fusiform, deep, robust, moderately laterally compressed. Dorsally dark gray; laterally silver with numerous, small, dark, irregular blotches; ventrally silver to white; dorsal part of head gray; fins dark gray. Head large, scaleless. Snout short. Eye placed laterally, on lower anterior end of head. Mouth large, terminal; lower jaw protrudes far forward and beyond upper jaw; upper jaw does not extend backward to anterior end of eye. Barbels absent. Lips fleshy. Teeth absent on jaws. Pharyngeal teeth long, 0,4-4,0. Gill rakers slender, long. Dorsal fin with 1 smooth spinous ray and 8-10 rays. Adipose fin absent. Caudal peduncle thick, slightly elongate. Caudal fin forked.

Anal fin falcate with 1 smooth spinous ray and 13-14 soft rays. Pelvic fins abdominal with 7-9 rays; insertion anterior to insertion of dorsal fin. Keel smooth, extending from pelvic fins to vent. Pectoral fins elongate, extending past insertion of pelvic fins; 1 spinous ray and 16-21 soft rays. Lateral line decurved and complete with 95-120 small cycloid scales in series. Adult males with sharp ridges on several anterior pectoral fin rays. Juveniles more silver dorsally and lack spinous rays in dorsal, anal, and pectoral fins.

Similar Species: Closely resembles the Silver Carp. Silver Carp solid silver on lateral sides without any dark, irregular blotches, and display a well-developed ventral keel that extends from the gills to the vent. Silver Carp also have pectoral fins that only extend to the insertion of the pelvic fins. Juvenile Bighead Carp may be mistaken for Golden 224

Shiner, which have smaller and fewer (42-54) scales in lateral line series. Grass Carp and

Common Carp have much larger cycloid scales.

Distribution and Habitat: Native to eastern China, eastern Siberia, and extreme eastern

North Korea, but have been extensively introduced worldwide.9 First introduced to the lower Mississippi River basin in United States in 1973 by the aquaculture industry as means of a food source, biological control of plankton in catfish aquaculture ponds, and water quality improvement in sewage treatment ponds.6,9,11,17 The species subsequently escaped through flooding or accidental releases and began to migrate up the tributaries of the Mississippi River, where it has since established itself in the Mississippi, Missouri,

Ohio, Illinois, and Tennessee River basins.9,17 Entered South Dakota through the

Missouri River in the late 1990’s, and now inhabits the Missouri River below Gavins

Point Dam, and the lower portions of the James, Vermillion and Big Sioux rivers in

South Dakota.9,17 Absent from North Dakota. Dispersion into these Missouri River watersheds has decreased or halted due to artificial and natural barriers.5 Primarily a large river species, strongly associated with areas >3 m (9.8 ft) in depth with low velocity (≤0.3 m/s) such as behind spur and wing dikes, but also inhabit lakes, reservoirs, and ponds.9 In the Illinois River, water temperature was negatively correlated with movement, suggesting that the species moves less when their growth optimum temperature of 26 °C

(78.8 °F) is exceeded.3 Juveniles often associated with backwaters rather than main channels of large rivers.9 Tolerant of extremes in water temperature. Preferred water temperature of young 25.0-26.9 °C (77-80.42 °F), with a critical thermal maximum of

38.8 °C (101.84 °F).9,2 In the Missouri River, tributary habitat use is highest in the winter, 225 with activity slowing at temperatures <4 °C (39.2 °F), and little movement occurring <2

°C (35.6 °F).9

Reproduction: Bighead Carp may migrate more than 80 km (49.71 mi) upstream to spawning grounds, which consist of rapid and turbid areas often near the confluence of rivers among rocks or sandbars.8,9 Spawning takes place during late summer and early fall in the United States.9 Spawning in the Missouri River was found to be initiated with increased water discharge and a water temperature of 22 °C (71.6 °F).9,16 Sexual maturity occurs at least by age-3 in the Missouri River.14 Prolific spawners. Spawning act takes place near the water’s surface and consists of two or more males actively chasing and head-butting females.9 Fecundity increases with size and age of female. In the lower

Missouri River, fecundity ranged 11,588-769,964 eggs, with an average of 226,213 eggs per female.14 No nest is constructed, and no parental care is given. Majority of eggs 0.5-

1.2 mm (0.02-0.05 in) in diameter, but may reach 1.5 mm (0.06 in) in diameter in the

Missouri River.14 Eggs semibuoyant and are carried with the current suspended in the water column until hatching.9 Hatching takes place in 24 hours at 28 °C (82.4 °F), and

30-32 hours at 25 °C (77 °F).1,9

Age and Growth: Growth rates in the Missouri River rather fast, with growth increments peaking between age-2 and age-3, and decreasing at ages >3.14 Age-3 individuals average

550 mm (21.65 in) in length, and age-5 individuals average 700 mm (27.56 in).14 Capable of reaching 1100 mm (43.31 in) TL and 18.14 kg (40 lbs) in the United States.7,9,14

Longevity 10 years in the Mississippi River.4

Food and Feeding: Larvae, juveniles, and adults opportunistic planktivores. Filter large amounts of plankton, specifically zooplankton, through gill rakers. Have been classified 226 as both “pump feeders”, using the buccal pump to push water through the gill rakers, and

“ram suspension feeders”, where the mouth is held open while swimming, forcing water through the gill rakers.9,10,13 Feeds near the surface during the evening and night within a wide range of water temperatures, and have been observed with full stomachs in the

Missouri River in water temperatures as low as 2.5 °C (36.5 °F).9 When zooplankton abundance is low, Bighead Carp will switch their diet to phytoplankton.9 Feeding habits of Bighead Carp often influence the size structure of plankton communities, and have the potential to negatively impact the growth of native fish species such as Paddlefish,

Gizzard Shad and Bigmouth Buffalo which also heavily depend on plankton.12,15

Literature Cited:

1. Bardach, J.E., J.H. Ryther, and W.O. McLarney. 1972. Aquaculture- the farming

and husbandry of freshwater and marine organisms. Wiley-Interscience, New

York.

2. Bettoli, P.W., W.H. Neill, and S.W. Kelsch. 1985. Temperature preference and

heat resistance of grass carp Ctenopharyngodon idella (Valenciennes), bighead

carp Hypophthalmichthys nobilis (Gray), and their F1 hybrid. Journal of Fish

Biology 27:239-247.

3. DeGrandchamp, K.L., J.E. Garvey, and R.E. Colombo. 2008. Movement and

habitat selection by invasive Asian carps in a large river. Transactions of the

American Fisheries Society 137:45-56.

4. Garvey, J.E., K.L. DeGrandchamp, and C.J. Williamson. 2006. Life history

attributes of Asian carps in the upper Mississippi River system. ANSRP Technical 227

Notes Collection (ERDC/ EL ANSRP-07-1), US Army Corps of Engineers

Research and Development Center, Vicksburg, Mississippi.

5. Hayer, C.A., B.D.S. Graeb, and K.N. Bertrand. 2014. Adult, juvenile and young-

of-year bighead, Hypophthalmichthys nobilis (Richardson, 1845) and silver carp,

H. molitrix (Valenciennes, 1844) range expansion on the northwestern front of the

invasion in North America. Biological Invasions Records 3:283-289.

6. Henderson, S. 1976. Observations on the bighead and silver carp and their

possible application in pond fish culture. Arkansas Game and Fish Commission,

Little Rock.

7. Howells, R.G. 2001. Introduced non-native fishes and shellfishes in Texas waters:

an updated list and discussion. Texas Parks and Wildlife Department

Management Data Series 188.

8. Jennings, D.P. 1988. Bighead carp (Hypophthalmichthys nobilis): a biological

synopsis. U.S. Fish and Wildlife Service, Washington, D.C. U.S. Fish and

Wildlife Service Biological Report 88:1-47.

9. Kolar, C.S., D.C. Chapman, W.R. Courtenay, Jr., C.M. Housel, J.D. Williams,

and D.P. Jennings. 2007. Bighead carps: a biological synopsis and environmental

risk assessment. American Fisheries Society, Special Publication 33, Bethesda,

Maryland.

10. Lu, M., and P. Xie. 2001. Impacts of filter-feeding fishes on the long-term

changes in crustacean zooplankton in a eutrophic subtropical Chinese lake.

Journal of Freshwater Ecology 16:219-228. 228

11. O’Connell, M.T., A.U. O’Connell, and V.A. Barko. 2011. Occurrence and

predicted dispersal of bighead carp in the Mississippi River system: development

of a heuristic tool. Pages 51-72 in D.C. Chapman and M.H. Hoff, editors. Invasive

Asian carps in North America. American Fisheries Society, Symposium 74,

Bethesda, Maryland.

12. Sampson, S.J., J.H. Chick, and M.A. Pegg. 2009. Diet overlap among two Asian

carp and three native fishes in backwater lakes on the Illinois and Mississippi

river. Biological Invasions 11:483-496.

13. Sanderson, S.L., J.J. Cech, Jr., and A.Y. Cheer. 1994. Paddlefish buccal flow

velocity during ram suspension feeding and ram ventilation. Journal of

Experimental Biology 186:145-156.

14. Schrank, S.J., and C.S. Guy. 2002. Age, growth, and gonadal characteristics of

adult bighead carp, Hypophthalmichthys nobilis, in the lower Missouri River.

Environmental Biology of Fishes 64:443-450.

15. Schrank, S.J., C.S. Guy, and J.F. Fairchild. 2003. Competitive Interactions

between age-0 bighead carp and paddlefish. Transactions of the American

Fisheries Society 132:1222-1228.

16. Schrank, S.J., P.J. Braaten, and C.S. Guy. 2001. Spatiotemporal variation in

density of larval bighead carp in the lower Missouri River. Transactions of the

American Fisheries Society 130:809-814.

17. Shearer, J.S. 2007. Exotic Species. Pages 253-266 in C. Berry, K. Higgins, D.

Willis, and S. Chipps, editors. History of fisheries and fishing in South Dakota.

South Dakota Department of Game, Fish and Parks, Pierre. 229

Common Shiner, Luxilus cornutus (Mitchill, 1817)

Etymology and Synonyms: Luxilus = Latin for luxilus, stemmed from Lux, meaning

“light”; cornutus = Latin for “horned”.

Description: Body fusiform, deep and laterally compressed. Dorsally dusky gray to olive with wide, prominent, dark mid-dorsal stripe present; laterally silver with lateral stripe fading anteriorly to dorsal fin; adults sometimes with small bits of concentrated pigment on some lateral scales to appear missing; ventrally silver to white; fins clear to lightly pigmented without any distinct markings. Head large. Snout bluntly pointed. Eye moderately large, placed laterally on head. Mouth large, terminal, oblique; upper jaw extending posteriorly between nostril and anterior edge of eye. Frenum absent. Barbels absent. Pharyngeal tooth pattern 2,4-4,2. Gill rakers short, roughly 9. Dorsal fin with 8 rays, straight distal end and rounded tip. Adipose fin absent. Caudal peduncle slender, elongate. Caudal fin moderately forked. Anal fin with 9 rays. Pelvic fins abdominal with

8 rays; insertions directly inferior or slightly posterior to insertion of dorsal fin. Pectoral fins with 15-17 rays. Lateral line complete, slightly decurved with 36-41 cycloid scales in series. Scales on anterior lateral side of body nearly 3 times greater in depth than width, and posterior edge of scale appearing rounded. Spawning males with faint, steel to iridescent blue on head and dorsal side of body; lateral sides of body and fins with rosy tint. Males also develop large tubercles on the snout and dorsal side of head, as well as on the nape and anterior rays of the dorsal and pectoral fins. Juveniles similar to adults with a decreased body depth.

Similar Species: Resembles the Golden Shiner, Red Shiner, Spotfin Shiner, and the

Rudd. Golden Shiner have a small, flattened, triangular shaped head, an elongate and 230 concave anal fin with 11-14 rays, and a lateral line with 42-54 cycloid scales in series.

Red Shiner are generally smaller, have a smaller eye, a shorter and thicker caudal peduncle, and spawning males with iridescent blue hue on body and red coloration on caudal, anal, pelvic and pectoral fins. Spotfin Shiner have a moderately deep body, and dark coloration forming a spot on the second or third posterior rays on the dorsal fin.

Rudd have a prominent, scaled keel anterior to the vent, 11-13 anal fin rays, and red to rosy coloration on all fins.

Distribution and Habitat: Native to North America throughout the upper Atlantic slope,

Great Lakes, the southern Hudson Bay, lower Missouri and upper Mississippi River basins.8 Present within the Missouri River, James, Sheyenne, Souris and Red River basins of North Dakota.10 More abundant and primarily found east of the Missouri River in

South Dakota within all the principle drainages, but have also been recorded from the

Niobrara, White, and Bad River drainages.1,4 Inhabits small to medium-sized streams of high gradient with pool and riffle sequences and clear to slightly turbid, cool water with sand, gravel or rubble substrate.2,5,11 May also occur in lakes and reservoirs, although less likely in the Dakotas.2,4 Often found schooling in the midwaters.11 Individuals from

Missouri streams had a reported mean critical dissolved oxygen concentration of 0.97 mg/L, and a mean critical thermal maximum as 35.7 °C (96.26 °F).13

Reproduction: Spawning takes place mid-May through late July, or when water temperatures reach 15.5-25.5 °C (59.9-77.9 °F).2,11,12,14 Males reach sexual maturity at roughly 77-120 mm (3.03-4.72 in) SL, and females at 75-120 mm (2.95-4.72 in) SL.9

Males prefer to utilize nests constructed by other species such as the Creek Chub, Central

Stoneroller or the Hornyhead Chub.2,12 If none of the preferred nest building species are 231 within the area, male Common Shiner will resort to constructing their own nest, which consists of them excavating a small depression with their snouts within shallow riffles over gravel substrate.2,12 Males defend small territories around the nest by chasing or attacking another male intruder.2,11 Spawning behavior consists of the male wrapping his caudal peduncle around the female and clasping onto her with his pectoral fins, which initiates the release of eggs and milt.2 Females are known to prefer spawning in nests in slower currents rather than faster currents.2 Eggs orange, demersal, adhesive to the bottom of the nest, and roughly 1.2-1.5 mm (0.05-0.06 in) in diameter.2

Age and Growth: Males generally experience faster growth rates than females.6,7 Mean lengths-at-age of males from the Des Moines River in Iowa are reported as: age-1, 46.34 mm (1.82 in) SL; age-2, 66.95 mm (2.64 in) SL; age-3, 78.25 mm (3.08 in) SL; age-4,

97.00 mm (3.82 in) SL; age-5, 116 mm (4.57 in) SL.6 Mean lengths-at-age of females are reported as: age-1, 42.12 mm (1.66 in) SL; age-2, 61.20 mm (2.41 in) SL.6 Capable of reaching 208.28 mm (8.2 in).15 Longevity 9 years, although they rarely live past age-6.2,16

Food and Feeding: Generalized, opportunistic omnivore. Feeding takes place either at the surface or in the midwater.11 Diets of adults and juveniles known to be relatively similar.3,6 Diets of individuals from Iowa were primarily made up of aquatic and terrestrial insects, but also consisted of filamentous algae, bottom ooze, small crustaceans, annelids, and fish.6,14 Water levels are known to greatly influence feeding.6

In Iowa, individuals were known to feed on greater amounts of plant matter during a flood period than animal matter.6

Literature Cited: 232

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Breder, C.M., and D.R. Crawford. 1922. The food of certain minnows. Zoologica

2:287-327.

4. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

5. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

6. Fee, E. 1965. Life history of the northern common shiner, Notropis cornutus

frontalis, in Boone County, Iowa. Proceedings of the Iowa Academy of Science

72:272-281.

7. Hubbs, C.L., and G.P. Cooper. 1936. Minnows of Michigan. Cranbrook institute

Science Bulletin 8:1-95.

8. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

9. Marshall, N. 1939. Annulus formation in scales of the common shiner, Notropis

cornutus chrysocephalus (Rafinesque). Copeia 1939:148-154.

10. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks. 233

11. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

12. Raney, E.C. 1940. The breeding behavior of the common shiner, Notropis

cornutus (Mitchill). Zoologica 25:1-14.

13. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

14. Starrett, W.C. 1950. Food relationships of the minnows of the Des Moines River,

Iowa. Ecology 31:216-233.

15. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

16. Van Oosten, J. 1932. The maximum age of fresh-water fishes. The Fisherman 1:3-

4.

234

Sturgeon Chub, Macrhybopsis gelida (Girard, 1856)

Etymology and Synonyms: Macr = “long” or “large”, hybopsis = genus of barbeled minnows; gelida = “frozen” or “stiff”.

Description: Body slender, elongate, and streamlined. Dorsally tan to light bronze; laterally silver with no distinct markings; silver to white; fins clear, transparent, lower lobe of caudal fin dusky gray with white edge. Head large, elongate. Snout elongate, fleshy, protruding well beyond mouth. Eye moderate, placed laterally on upper portion of head. Mouth large, subterminal. Frenum absent. Barbels present, one on each corner of mouth; slender, prominent. Lips thin. Pharyngeal tooth pattern 1,4-4,1. Gill rakers blunt,

3-6. Dorsal fin with 8 rays, straight distal end and rounded tip. Adipose fin absent.

Caudal peduncle slender, elongate. Caudal fin forked. Anal fin with 8 rays, straight distal end and rounded tip. Pelvic fins abdominal and rounded with 8 rays; insertion over or slightly anterior of dorsal fin insertion. Pectoral fins with 15-17 rays; do not extend beyond insertions of pelvic fins when depressed. Lateral line complete with 39-45 cycloid scales in series. Scales on dorsal side with a longitudinal ridge or keel present.

Spawning males develop small tubercles on pectoral fins. Juveniles similar in appearance to adults.

Similar Species: Closely resembles the Sicklefin Chub. Sicklefin Chub lack any longitudinal ridges or keels on dorsal scales, and have sickle shaped dorsal and pectoral fins. Pectoral fins of the Sicklefin Chub are also elongate, and extend well beyond the insertion of the pelvic fins when depressed. Easily distinguished from Flathead Chub,

Silver Chub, Plains Minnow and Western Silvery Minnow by the presence of longitudinal ridges or keels on dorsal scales. Shoal Chub display small, dark specks on 235 dorsal and lateral sides. May also be mistaken for Longnose Dace but can be distinguished by the lack of a frenum.

Distribution and Habitat: Native to the mainstem and many tributaries of the Missouri

River from Montana and Wyoming in the northwest, south through the lower Mississippi

River to Louisiana and the Gulf of Mexico.12 Occurs throughout the Little Missouri,

Yellowstone, and Missouri rivers in North Dakota.2,13 Inhabits isolated potions of the

Missouri River and its western tributaries including the White and Cheyenne rivers of

South Dakota.1,3 Extant populations have been reduced by more than 70% in their native range likely due to the construction of reservoirs and dewatering.3,6,7,9 Because the species requires long free-flowing sections of river, abundance is reduced downstream of impoundments for an estimated minimum threshold of 297 km (184.5 mi).6 Preferred habitat includes the benthic zone of medium to large turbid rivers with sand, rock, or gravel riffle zones with an average depth of 0.23 m (0.75 ft) and moderate to fast current with an average velocity of 0.55 m/s.7,10,12 In the upper Missouri and lower Yellowstone rivers in western North Dakota, the species was collected most frequently from main channels, but also occurred within sandbars and border channels.2,13 Presence of Sturgeon

Chub also increases with decreasing depth, increasing velocity and decreasing water clarity.2 Adapted to and tolerant of highly turbid waters.12 Juveniles have been known to occupy shallow areas approximately 0.6 m (1.97 ft) in depth with sandy substrate, moderate velocity and no aquatic vegetation, however it has also been suggested that they do not require different habitat than mature adults.2,10 Rarely occurs in backwater habitat.2

Reproduction: Information regarding the spawning and reproductive behavior of

Sturgeon Chub is limited. Spawning occurs mid-June through July when water 236 temperatures reach 18.3-22.2 °C (64.94-71.96 °F).10,4,14 Pelagic, broadcast and fractional spawners.3,8,10 Sexual maturity occurs at age-2, or when females reach approximately 76-

81 mm (2.99-3.19 in) TL, and males 78-79 mm (3.07-3.11 in) TL.10,14 No nest is constructed and no parental care is given. Fecundity ranges roughly 2,000-3,500 eggs per female.10 Eggs are carried with the current downstream during incubation and larval development.3,11 Reproductive success or the survival of young-of-the-year is suspected to be low.10

Age and Growth: Mean lengths-at-age of individuals from the upper Missouri and lower

Yellowstone rivers in western North Dakota are reported as: age-1, 42.5 mm (1.67 in)

TL; age-2, 65.4 mm (2.57 in) TL; age-3, 82.8 mm (3.26 in) TL.2 Capable of reaching

96.5 mm (3.80 in) TL.10 Longevity 4 years.10

Food and Feeding: Benthic insectivore. Information on diet composition is scarce.

Adults are known to feed on small aquatic invertebrates living within the rocky substrate in riffle zones.10 Age-0 individuals primarily feed on midge larvae such as chironomids.8

Sturgeon Chub utilize their highly developed cutaneous sensory system including maxillary barbels and numerous external taste buds on the ventral side of the head and body to forage in highly turbid waters.5,10

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Everett, S.R., D.L. Scarnecchia, and L.F. Ryckman. 2004. Distribution and habitat

use of sturgeon chubs (Macrhybopsis gelida) and sicklefin chubs (M. meeki) in 237

the Missouri and Yellowstone Rivers, North Dakota. Hydrobiologia 527: 183-

193.

3. Hoagstrom C.W., C.A. Hayer, J.G. Kral, S.S. Wall, and C.R. Berry Jr. 2006. Rare

and declining fishes of South Dakota: a river drainage scale perspective.

Proceedings of the South Dakota Academy of Science 85:171-211.

4. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

5. Moore, G.A. 1950. The cutaneous sense organs of barbeled minnows adapted to

life in the muddy waters of the Great Plains region. Transactions of the American

Microscopy Association 69:69-95.

6. Perkin, J.S., and K.B. Gido. 2011. Stream fragmentation thresholds for a

reproductive guild of Great Plains fishes. Fisheries 36:371-383.

7. Reigh, R.C., and D.S. Elsen. 1979. Status of the sturgeon chub (Hybopsis gelida)

and sicklefin chub (H. meeki) in North Dakota. The Prairie Naturalist 11:49-52.

8. Starks, T.A., M.L. Miller, and J.M. Long. 2016. Early life history of three pelagic-

spawning minnows Macrhybopsis spp. in the lower Missouri River. Journal of

Fish Biology 88:1335-1349.

9. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery 238

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67.

10. Stewart, D.D. 1981. The biology of the sturgeon chub (Hybopsis gelida Girard) in

Wyoming. M.S. Thesis, University of Wyoming, Laramie.

11. Tibbs, J.E., and D.L. Galat. 1997. Larval, juvenile, and adult small fish use of

scour basins connected to the lower Missouri River. Report to the Missouri

department of Conservation, Jefferson City.

12. U.S. Fish and Wildlife Service. 1993. Status report on sturgeon chub

(Macrhybopsis gelida), a candidate endangered species. U.S. Fish and Wildlife

Service, Bismarck, North Dakota.

13. Welker, T.L., and D.L. Scarnecchia. 2004. Habitat use and population structure of

four native minnows (family Cyprinidae) in the upper Missouri and lower

Yellowstone rivers, North Dakota (USA). Ecology of Freshwater Fish 13:8-22.

14. Werdon, S.J. 1992. Population status and characteristics of Macrhybopsis gelida,

Platygobio gracilis, and Rhinichthys cataractae in the Missouri River basin. MS

thesis, South Dakota State University, Brookings, SD.

239

Shoal Chub, Macrhybopsis hyostoma (Gilbert, 1884)

Etymology and Synonyms: Macr = “long” or “large”, hybopsis = genus of barbeled minnows; hyo = Greek for “hog”, stoma = “mouth”.

Description: Body fusiform, slender, elongate, slightly dorsoventrally flattened. Dorsally tan to bronze with small, prominent, black, scattered specks; laterally silver with small, prominent, black, scattered specks; ventrally silver to white; fins clear with no distinct markings. Head elongate. Snout fleshy, slightly elongate, extending well beyond mouth.

Eye large, placed laterally on head. Mouth small, subterminal. Frenum absent. Barbels present, one, sometimes two on each corner of mouth; small and conspicuous. Lips thin, fleshy. Pharyngeal tooth pattern 0,4-4,0. Dorsal fin with 8 rays, nearly straight distal end and rounded tip. Adipose fin absent. Caudal peduncle slender, elongate. Caudal fin forked. Anal fin with 7-8 rays, straight distal end and rounded tip. Pelvic fins abdominal, short and rounded; insertion directly under or slightly anterior to insertion of dorsal fin.

Pectoral fins large, rounded to slightly pointed; do not extend beyond insertion of pelvic fins when depressed. Lateral line complete with 32-43 cycloid scales in series. Scales typically absent on breast. Spawning males develop small tubercles on dorsal sides of head and body, as well as the breast, and rays of the pectoral and pelvic fins. Juveniles similar in appearance to adults.

Similar Species: Easily distinguished from Sicklefin and Sturgeon Chub by the large eye and small, prominent, black, scattered specks on the dorsal and lateral sides. Sicklefin

Chub also display sickle shaped and elongate pectoral fins that extend well beyond the insertion of the pelvic fins. Sturgeon Chub display longitudinal ridges or keels on the dorsal scales. Flathead Chub have a dorsoventrally flattened head and lack dark specks on 240 body. Silver Chub have a slightly laterally compressed body lacking any dark specks on dorsal and lateral sides.

Distribution and Habitat: Native throughout much of the south-central United States and Mississippi River Basin from southeastern South Dakota, southern Minnesota and

Wisconsin in the north, south to Texas, and from the Appalachian Mountains in the east to the Rio Grande River in the west.8 Absent from North Dakota. Occurs in South Dakota throughout the Missouri River below Gavins Point Dam. Extant populations throughout the native range have been experiencing significant declines, likely due to anthropogenic disturbances such as river channelization and impoundments.6,11 Inhabits the benthic zone within tributaries and mainstems of small to large, turbid rivers with moderate to strong current over medium to large gravel riffles and raceways.2,5 In the upper Mississippi

River Basin, Shoal Chubs preferred water temperatures of 158-179.6 °C (70-82 °F) with moderate current ranging 0.38-0.57 m/s, and relatively shallow depths ranging 1.04-2.69 m (3.41-8.83 ft).5 Known to occupy shallow waters during night.2,7 Juveniles prefer areas with sand substrate.5

Reproduction: Information regarding the spawning and reproductive behavior of Shoal

Chub is limited, but they are likely one of the several species of pelagic, broadcast spawners in the Great Plains region.9,12 Spawning is prolonged, and takes place from early spring until late fall.1,4,12 Sexual maturity occurs at age-1.12 Mean clutch size from mature females 39-70 mm (1.54-2.76 in) TL was 198.75, and ranged from 34-680 oocytes per female.12 Clutch size is positively related to the total length of mature females.12 Fertilized eggs are roughly 2.5 mm (0.10 in) in diameter, transparent, semibuoyant, nonadhesive, and carried with the current downstream during incubation 241 and larval development.2,9,10 Although river fragmentation created by impoundments negatively impacts the upstream migration of Shoal Chub, it is suggested that the species requires relatively shorter unsegmented stream lengths for the completion of their life history than other pelagic-spawning cyprinids.9,10

Age and Growth: Growth known to be rapid during the first year of life, with individuals in Texas reported reaching 45-65% of their maximum length during the first summer and fall.12 Capable of reaching 76 mm (2.99 in) TL.4,8 Longevity 2.5 years, with little amount of individuals surviving to age-2.12

Food and Feeding: Benthic insectivore.5 Adults primarily consume small aquatic invertebrates such as chironomids, dipterans, and trichoptera larvae.5,12 Juveniles consume detritus, sand, aquatic insects and small crustaceans.12 Shoal Chub differ than other species of the genus Macrhybopsis, in that they are primarily a sight feeder given their larger eye, patterns in brain morphology and distribution of external taste buds.3

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Davis, B.J., and R.J. Miller. 1967. Brain patterns in minnows of the genus

Hybopsis in relation to feeding habits and habitat. Copeia 1967:1-39.

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville. 242

5. Gaughan, S. 2016. Habitat use, molecular phylogeny, and population structure of

Macrhybopsis chubs in the upper Mississippi River basin. M.S. Thesis, University

of Nebraska.

6. Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, 5.

Selected chubs and minnows (Cyprinidae): sicklefin chub (M. meeki), sturgeon

chub (M. gelida), silver chub (M. storeriana), speckled chub (M. aestivaus),

flathead chub (Platygobio gracilis), plains minnow (Hybognathus placitus), and

western silvery minnow (H. argyritis). Transactions of the Nebraska Academy of

Sciences and Affiliated Societies 21:99-108.

7. Klutho, M.A. 1983. Seasonal, daily, and spatial variation of shoreline fishes in the

Mississippi River at Grand Tower, Illinois. M.S. Thesis, Southern Illinois

University, Carbondale, Illinois.

8. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

9. Perkin, J.S., and K.B. Gido. 2011. Stream fragmentation thresholds for a

reproductive guild of Great Plains fishes. Fisheries 36:371-383.

10. Platania, S.P., and C.S. Altenbach. 1998. Reproductive strategies and egg types of

seven Rio Grande Basin cyprinids. Copeia 1998:559-569.

11. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery 243

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67.

12. Williams, C.S. 2011. Life history characteristics of three obligate riverine species

and drift patterns of Lower Brazos River fishes. Ph.D. Dissertation, Texas State

University, San Marcos, Texas.

244

Sicklefin Chub, Macrhybopsis meeki (Jordan & Evermann, 1896)

Etymology and Synonyms: Macr = “long” or “large”, hybopsis = genus of barbeled minnows; meeki = named after Seth E. Meek, an early American ichthyologist in central

North America.

Description: Sicklefin Chub have a slender, elongate and streamlined body. Dorsally tan to light bronze; laterally silver with no distinct markings; ventrally silver to white; fins clear, transparent, except lower lobe of caudal fin which is dusky gray with white edge.

Head large and blunt. Snout large and blunt; protrudes past the upper jaw. Eye moderate, placed laterally on upper portion of head. Mouth large, subterminal. Barbels are short and prominent with one on each corner of mouth. Lips thin. Pharyngeal tooth pattern 0,4-4,0.

Gill rakers blunt; 5. Dorsal fin is sickle shaped with pointed tip and 8 rays; insertion over or slightly posterior of pelvic fin insertion. Adipose fin absent. Caudal peduncle elongate, relatively thin, uniform in thickness. Caudal fin forked. Anal fin has 8 rays and a pointed tip. Pelvic fins abdominal, sickle shaped and pointed. Pectoral fins are long and sickle shaped with 16-17 rays; extend beyond insertion of pelvic fins when depressed. Lateral line complete with 43-50 cycloid scales in series. Scales on dorsal side lack a longitudinal ridge or keel. No nuptial coloration on body or fins, however small tubercles develop on all fins except the caudal fin.

Similar Species: Closely resembles the Sturgeon Chub. Sturgeon Chub have a longitudinal ridge or keel present on each of the dorsal scales. Flathead Chub have a broad, dorsoventrally flattened head with a snout tapering into a point with a rounded tip, and pectoral fins that do not extend past the insertion of the pelvic fins when depressed. 245

Silver Chub, Plains Minnow and Western Silvery Minnow have larger eyes, and pectoral fins that do not extend past the insertion of the pelvic fins when depressed.

Distribution and Habitat: Primarily found within the main stem and tributaries in the

Missouri and lower Yellowstone Rivers of Montana, and south to the middle Mississippi

River from the Missouri River’s confluence to the Ohio Rivers confluence.4,14 The distribution and abundance of Sicklefin Chub have declined throughout much of the species historical range, likely due to impoundments and channelization which reduce turbidity, alter the flow regimes, and reduce early life stage habitat.13,15,19 The species requires long, free flowing reaches with a natural range of variability in seasonal low flows and turbidity.4 In the upper Missouri and lower Yellowstone River segments of western North Dakota, Sicklefin Chubs were collected most frequently from the main channels, but also occurred within sandbars, border and side channels.7 Habitat use changes throughout the Sicklefin Chubs lifetime.13 Post hatching, larvae swim vertically in the water column and rarely rest along the bottom.1 Moderately shallow (average 2.1 m

(6.89 ft.)), riverine shoreline areas with relatively slow velocities (mean 0.30 m/sec) provide critical nursery habitat.4,13 Adults inhabit areas with higher water velocities

(mean 0.93 m/sec), and moderate depths (average 3.1 m (10.17 ft.)).13 The presence of

Sicklefin Chub increases with increased water depth, decreasing velocity and decreasing water clarity.7 Sicklefin Chub rarely occur in reservoirs and backwaters.8,7,10,20

Reproduction: Spawning in the lower Missouri River is protracted, beginning in early

June, peaking in mid-July, and subsiding in August.18 Minimum spawning water temperature 20°C (68°F).2 Sicklefin Chub are pelagic, broadcast, and fractional spawners.1 Sexual maturity occurs at age-3.5 Spawning known to take place at night.1 246

Spawning behavior consists of one or two males aligning their bodies laterally with a single female in a head-to-head orientation, and nudging her to swim in a circular motion near the substrate.1 While swimming in a circular motion, the couple will embrace, with the male applying pressure and wrapping his caudal fin and caudal peduncle around the female, initiating the release of eggs and milt.1 No nest is constructed and no parental care is given.1 In aquaria, 104-577 eggs were released during a single spawning event, but females are capable of producing at least 1,561 eggs.1,5 Water hardened eggs are on average 3.41 mm (0.13 in) in diameter, semi-buoyant, non-adhesive, and are carried with the current downstream during incubation and larval development.1,16 Eggs may sink if water current is not maintained at an appropriate level, decreasing survival rates.1,16

Sicklefin Chub likely exhibit a high degree of post-spawning mortality.5

Age and Growth: Larvae average 5.19 mm (0.20 in) at hatching.1 Minimum growth water temperature 10°C (50°F).2 First year growth rates are significantly greater in higher latitudes than low latitudes.2 Mean lengths-at-age of individuals from the upper Missouri and lower Yellowstone rivers in western North Dakota are reported as: age-1, 46.0 mm

(1.81 in) TL; age-2, 75.3 mm (2.96 in) TL; age-3, 93.1 mm (3.67 in) TL; age-4, 107 mm

(4.21 in) TL.7 Maximum age is also known to be significantly related to latitude and increases from low to high latitudes.2 Capable of reaching 128 mm (5.04 in) TL.6

Longevity 4 years.9,19

Food and Feeding: Benthic insectivores.20 Adult diets primarily consist of small, benthic, adult and larval aquatic insects including Dipterans, Ephemeropterans, and

Trichopterans.12 Sicklefin Chub have developed unique adaptations to forage in highly 247 turbid waters including elaborate internal and external taste buds, lateral line neuromasts, and numerous small sensory papillae.3,4,11,17

Literature Cited:

1. Albers, J.L., and M.L. Wildhaber. 2017. Reproductive strategy, spawning

induction, spawning temperatures and early life history of captive sicklefin chub

Macrhybopsis meeki. Journal of Fish Biology 91:58-79.

2. Braaten, P.J., and C.S. Guy. 2002. Life history attributes of fishes along the

latitudinal gradient of the Missouri River. Transactions of the American Fisheries

Society 131:931-945.

3. Davis, B.J., and R.J. Miller. 1967. Brain patterns in minnows of the genus

Hybopsis in relation to feeding habits and habitat. Copeia 1967:1-39.

4. Dieterman, D.J., and D.L. Galat. 2004. Large-scale factors associated with

sicklefin chub distribution in the Missouri and lower Yellowstone rivers.

Transactions of the American Fisheries Society 133:577-587.

5. Dieterman, D.J., E. Roberts, P.J. Braaten, and D.L. Galat. 2006. Reproductive

development in the sicklefin chub, Macrhybopsis meeki, in the Missouri River

and Yellowstone River USA. Prairie Naturalist 38:113-130.

6. Dieterman, D.J., M.P. Ruggles, M.L. Wildhaber, and D.L. Galat. 1996.

Population structure and habitat use of benthic fishes along the Missouri and

lower Yellowstone rivers. 1996 Annual Report of the Missouri River Benthic Fish

Study PD-95-5832 to US Army Corps of Engineers and US Bureau of

Reclamation. 248

7. Everett, S.R., D.L. Scarnecchia, and L.F. Ryckman. 2004. Distribution and habitat

use of sturgeon chubs (Macrhybopsis gelida) and sicklefin chubs (M. meeki) in

the Missouri and Yellowstone Rivers, North Dakota. Hydrobiologia 527:183-193.

8. Fisher, S.J. 1999. Seasonal investigation of native fishes and their habitats in

Missouri River and Yellowstone River backwaters. Ph.D. Dissertation, South

Dakota State University, Brookings, South Dakota.

9. Herman, P., A. Plauck, N. Utrup, and T. Hill. 2008. Three year summary age and

growth report for sturgeon chub (Macrhybopsis gelida). Report to the US Army

Corps of Engineers, Northwestern Division, Omaha, Nebraska.

10. Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, 5.

Selected chubs and minnows (Cyprinidae): sicklefin chub (M. meeki), sturgeon

chub (M. gelida), silver chub (M. storeriana), speckled chub (M. aestivaus),

flathead chub (Platygobio gracilis), plains minnow (Hybognathus placitus), and

western silvery minnow (H. argyritis). Transactions of the Nebraska Academy of

Sciences and Affiliated Societies 21:99-108.

11. Moore, G.A. 1950. The cutaneous sense organs of barbeled minnows adapted to

life in the muddy waters of the Great Plains region. Transactions of the American

Microscopy Association 69:69-95.

12. Nocomis, E. 2014. Observations of sicklefin chub diets in the Missouri River. The

Prairie Naturalist 46:88-91.

13. Panella, M.J., S.C. Schainost, G.E. Mestl, and K.D. Steffensen. 2018. Listing

proposal for four small-bodied fishes in Nebraska: Flathead chub (Platygobio

gracilis), plains minnow (Hybognathus placitus), sicklefin chub (Macrhybopsis 249

meeki), and western silvery minnow (Hybognathus argyritis). Nebraska Game

and Parks Commission, Lincoln, Nebraska.

14. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of

Conservation, Jefferson City, Missouri.

15. Pflieger, W.L., and T.B. Grace. 1987. Changes in the fish fauna of the lower

Missouri River, 1940-1983. Pages 166-177 in Community and evolutionary

ecology of North American stream fishes. W. Matthews and D. Heins (editors).

University of Oklahoma Press, Norman.

16. Platania, S.P., and C.S. Altenbach. 1998. Reproductive strategies and egg types of

seven Rio Grande Basin cyprinids. Copeia 1998:559-569.

17. Reno, H.W. 1969. Cephalic lateral-line systems of the cyprinid genus Hybopsis.

Copeia 1969:736-773.

18. Starks, T.A., M.L. Miller, and J.M. Long. 2016. Early life history of three pelagic-

spawning minnows Macrhybopsis spp. in the lower Missouri River. Journal of

Fish Biology 88:1335-1349.

19. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67. 250

20. Werdon, S.J. 1993. Status report on sicklefin chub (Macrhybopsis meeki), a

candidate endangered species. U.S. Fish and Wildlife Service, Bismarck, North

Dakota.

251

Silver Chub, Macrhybopsis storeriana (Kirtland, 1845)

Etymology and Synonyms: Macr = “long” or “large”, hybopsis = genus of barbeled minnows; storeriana = named after David H. Storer, an early North American ichthyologist.

Description: Body fusiform, slightly stout and laterally compressed body. Dorsally tan to dark gray; laterally silver with no distinct markings; ventrally silver to white. Head short.

Snout blunt, rounded and fleshy protruding beyond mouth. Eye large, placed laterally on head. Mouth small, subterminal. Frenum absent. Barbels present, one at each corner of the mouth; short, conical, inconspicuous. Lips thin, fleshy. Pharyngeal tooth pattern 1,4-

4,1. Dorsal fin with 8 rays, nearly straight distal end and slightly pointed tip. Adipose fin absent. Caudal peduncle slender, elongate. Caudal fin forked. Anal fin with 7-8 rays, nearly straight distal end and pointed tip. Pelvic fins abdominal, short and rounded distal end with 8 rays; insertion slightly posterior to insertion of dorsal fin. Pectoral fins with

17-18 rays; do not extend beyond insertion of pelvic fins when depressed. Lateral line complete with 35-48 cycloid scales in series. Scales typically absent on breast, but occasionally present on posterior breast. Spawning males develop large tubercles on pectoral fin rays, as well as smaller tubercles on the dorsal and lateral sides of the head.

Juveniles similar in appearance to adults.

Similar Species: Closely resembles Shoal Chub, Sicklefin Chub and Sturgeon Chub.

Shoal Chub display small, prominent, black, scattered specks along the dorsal and lateral sides of the body. Sicklefin Chub have smaller eyes and sickle shaped, elongate pectoral fins that extend well beyond the insertion of the pelvic fins. Sturgeon Chub have smaller 252 eyes and display longitudinal ridges or keels on the dorsal scales. Flathead Chub have a dorsoventrally flattened head with smaller eyes, and a snout that tapers to a point.

Distribution and Habitat: Native throughout the Mississippi River basin from the

Missouri River in western Nebraska, east to the Monongahela River in Pennsylvania, and from the upper Mississippi River along the Minnesota-Wisconsin border, south to the lower Mississippi River in Lousiana.5,11,12,13,16 Lake Erie of the Great Lakes is known to support the only lake-dwelling population of Silver Chub.12 Disjunct populations occur within the Red River in North Dakota.12,13 Occurs within the Missouri River and a few watersheds within the Keya Paha, James, Big Sioux, and Niobrara river basins in South

Dakota.7,8 Macrohabitat generalists, inhabiting large, low gradient, semi-turbid bodies of water including lakes, impoundments, rivers and headwaters of tributaries with stronger currents.6,11,17 Rarely occurs within small streams, but occasionally inhabits headwaters of small streams.11 Occurs in areas with silt, clay, sand, and small gravel substrate.8,11,18 In the Mississippi River, overwintering habitat consists of deep holes.1

Reproduction: Information regarding the spawning and reproductive behavior of the

Silver Chub is limited, especially in lotic habitats, however they are likely one of the several species of pelagic, broadcast spawners in the Great Plains region.14 Spawning is suggested to take place late May through early June, or when water temperature reaches roughly 20 °C (68 °F). 1,2,6,11 Sexual maturity occurs at age-1.11 Seasonal migrations upstream have been documented, suggesting spawning migrations in springtime.3 No nest is constructed and no parental care is given. Fecundity generally increases with the size of female.11 Mature eggs light-orange in color, semibuoyant, and likely carried with the current downstream during incubation and larval development.11,14 Mature eggs measured 253

0.9 mm (0.04i n) in diameter form an age-1 female, and 1.7 mm (0.07 in) in diameter from an age-3 female.2 It is suggested that 203 km (126.14 mi) of unsegmented river are necessary for the Silver Chub to complete their life history and stabilize populations.14

Age and Growth: Information on age and growth of the species is scarce. Majority of growth takes place during summer months, with little to no growth occurring during the winter.11 Capable of reaching 231 mm (9.09 in) TL.13 Longevity 4 years, with females generally living longer than males.11

Food and Feeding: Generalist benthic feeders.17 Age-0 individuals from western Lake

Erie fed on microcrustacea such as copepods and Daphnia, as well as small immature insects such as chironomid larvae and pupae.11 Adults consume Ephemeroptera nymphs, such as Hexagenia., small mollusks, and Daphnia.5,11,12,15 At water temperatures 18-22 °C

(64.4-71.6 °F), it was found that adult Silver Chubs are able to consume between 4-5% of their body weight in food per day, and juveniles can consume roughly 10% of their body weight in food per day.11 Silver Chub use external taste buds on the head and pectoral fins to detect prey, but likely also utilize their larger eye and better sense of sight to assist in foraging, unlike Sturgeon or Sicklefin Chub.4,9,10

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357. 254

4. Davis, B.J., and R.J. Miller. 1967. Brain patterns in minnows of the genus

Hybopsis in relation to feeding habits and habitat. Copeia 1967:1-39.

5. Gaughan, S. 2016. Habitat use, molecular phylogeny, and population structure of

Macrhybopsis chubs in the upper Mississippi River Basin. M.S. Thesis,

University of Nebraska-Omaha, Nebraska.

6. Harlan, J.R., and E.B. Speaker. 1987. Iowa Fish and Fishing. Iowa Department of

Natural Resources, Des Moines, Iowa.

7. Harland, B., and C.R. Berry, Jr. 2004. Fishes and habitat characteristics of the

Kaya Paha River, South Dakota-Nebraska. Journal of Freshwater Ecology

19:169-177.

8. Hayer, C.A., B.C. Harland, and C.R. Berry, Jr. 2006. Recent range extensions,

name changes and status updates for selected South Dakota fishes. Proceedings of

the South Dakota Academy of Science 85:247-265.

9. Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, 5.

Selected chubs and minnows (Cyprinidae): sicklefin chub (M. meeki), sturgeon

chub (M. gelida), silver chub (M. storeriana), speckled chub (M. aestivaus),

flathead chub (Platygobio gracilis), plains minnow (Hybognathus placitus), and

western silvery minnow (H. argyritis). Transactions of the Nebraska Academy of

Sciences and Affiliated Societies 21:99-108.

10. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence. 255

11. Kinney, E.C. Jr., 1954. A life history study of the silver chub, Hybopsis

storeriana (Kirtland), in western Lake Erie with notes on associated species.

Ph.D. Dissertation, Ohio State University, Columbus, Ohio.

12. Kočovský, P.M. 2018. Diets of endangered silver chub (Macrhybopsis storeriana,

Kirtland, 1844) in Lake Erie and implications for recovery. Ecology of

Freshwater Fish 2018:1-8.

13. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

14. Perkin, J.S., and K.B. Gido. 2011. Stream fragmentation thresholds for a

reproductive guild of Great Plains fishes. Fisheries 36:371-383.

15. Ross, S.T. 2001. Inland fishes of Mississippi. University Press of Mississippi,

Jackson, Mississippi.

16. Stauffer, J.R. Jr., R.W. Criswell, and D.P. Fischer. 2016. The Fishes of

Pennsylvania. Cichlid Press, El Paso, Texas.

17. Steffensen, K.D., D.A. Shuman, and S. Stukel. 2014. The status of fishes in the

Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub

(M. gelida), sicklefin chub (M. meeki), silver chub (M. storeriana), flathead chub

(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery

minnow (H. argyritis), and brassy minnow (H. hankinsoni). Transactions of the

Nebraska Academy of Sciences and Affiliated Societies 34:49-67. 256

18. Wood, K.G. 1953. Distribution and ecology of certain bottom-living invertebrates

of the western basin of Lake Erie. Ph.D. Dissertation, Ohio State University,

Columbus, Ohio.

257

Northern Pearl Dace, Margariscus nachtriebi (Cox, 1896)

Etymology and Synonyms: Margariscus = Greek for “margariskos”, meaning “pearl”; nachtriebi = unknown.

Description: Body fusiform, stout, slightly laterally compressed posteriorly. Dorsally dusky brown to dark olive; laterally light bronze to olive, mottled with small irregular dark specks and single dark, dusky lateral stripe that is more prominent on posterior end of body, and fades with age and growth; ventrally cream to white; fins lightly pigmented.

Head moderate. Snout blunt. Eye moderately large, placed laterally on head. Mouth terminal, slightly oblique, relatively small; jaw extends to below nostril. Frenum absent.

Lips fleshy; groove on upper lip continuous over snout. Barbels sometimes present; small, flattened, one on each side of mouth between the lip and upper jaw. Pharyngeal tooth pattern 2,5-4,2. Gill rakers short, 6-8. Dorsal fin slightly rounded with 8 rays.

Adipose fin absent. Caudal peduncle elongate, moderately thick. Caudal fin moderately forked. Anal fin with 8 rays. Pelvic fins abdominal, slightly rounded with 8 rays; insertion distinctly anterior to insertion of dorsal fin. Pectoral fins slightly rounded with

15-16 rays. Lateral line complete with 60-75 small cycloid scales in series. Spawning males develop small tubercles on the head, pectoral fin rays, lateral sides and on breast.

Males also develop a prominent red-orange stripe on flank below the dark, dusky lateral stripe. Spawning females may also develop small tubercles on pectoral fin rays, as well as a golden yellow stripe on flank below the dark, dusky lateral stripe. Juveniles with a more distinct and prominent lateral stripe, ending in a dark spot at the base of the caudal fin.

Similar Species: Closely resembles the Creek Chub, Lake Chub and Finescale Dace.

Creek Chub have a dark spot present at the anterior base of the dorsal fin, as well as 258 barbels that are more conical in shape. Lake Chub are more gray to silver in color, have a slightly subterminal mouth extending past the nostril, and have longer barbels always present in the corners of the mouth. Finescale Dace display an incomplete lateral line, appear scaleless with the naked eye, and have an upper jaw that extends to anterior end or middle of eye.

Distribution and Habitat: Native throughout most of Canada and in the United States throughout the Missouri, Mississippi and Great Lakes-St. Lawrence River drainages from northern Montana and North Dakota in the west, east throughout the northern states to

Maine, and south to Virginia, with isolated populations in Iowa, South Dakota and

Nebraska.4,8,10,13,14 Occur within the Red River of the North, Souris, Missouri, Heart,

Knife, and Little Missouri River basins in North Dakota. Inhabit coolwater small glacial lakes, beaver ponds, bogs and creeks in the northern part of their range, but are more commonly found in the cool, springfed, headwater stream pools of perennial first and second order streams in South Dakota.4,8,12,15,16 The species occurrence in the Keya Paha,

White River, and Little White River basins in southwestern South Dakota and the

Nebraska Sandhills region is relict of Palistocene glaciaion.1,5,6 Preferred habitat in this region includes areas of meandering streams with well vegetated stream banks, abundant macrophyte growth, undercut banks, and the absence of large piscivores.4 Optimum water temperature roughly 16 °C (60.8 °F).4 Adults are generally restricted to the mid-water column and deeper pools, while juveniles often occupy areas higher in the water column closer to instream vegetation.3,4 However in winter months, all age classes reside in deep pools.3,17 Capable of tolerating a broad range of environmental conditions in perennial 259 headwater streams, which likely explains the species extensive range.8 Moderately intolerant of turbidity.18

Reproduction: Spawning takes place mid-April to mid-May, or when water temperatures reach 16-18 °C (60.8-64.4 °F).3,4 Spawning also dependent on photoperiod.4

Sexual maturity reached at age-1 in both sexes.16 Spawning behavior has only been described once from a Michigan stream where males were reported to coax gravid females into a small territory in which they defended.9 Males positioned themselves parallel to the female grasping onto her with their tuberculate pectoral fins and wrapped their caudal fin around her caudal peduncle.9 The pair then vibrated their bodies together for roughly 2 seconds to initiate the release of eggs and milt.9 Lithophilic spawner; deposits eggs over gravel or rocky substrate in areas with little to no current.4 No nest is constructed and no parental care is given.4,9 Clutch size in Nebraska is variable with age-1 females averaging 1,100 eggs and age-2 females averaging 2,800 eggs.3,4 Eggs roughly

1.24-1.40 mm (0.05-0.06 in) in diameter.2,7

Age and Growth: Mean standard lengths-at-age of females from Nebraska are reported as: age-1, 56.1 mm (2.21 in) SL; age-2, 68.5 mm (2.70 in) SL; age-3, 78.6 mm (3.09 in)

SL; age-4, 88.3 mm (3.48 in) SL.16 Females generally live longer, grow faster, and attain greater sizes than males.16 Capable of reaching 158 mm (6.22 in) TL.2,11 Longevity 4 years.16

Food and Feeding: Sight-feeding omnivores that actively forage throughout the day.4,17

Individuals <54 mm (2.13 in) TL primarily feed on Diptera larvae, algae, Copepods and

Cladocerans.16 Fish >74 mm (2.91 in) TL have a more diverse diet consisting of larger organisms such as snails, fingernail clams, amphipods and larger insects often found in 260 the benthic zone.16 Sand, silt and detritus are also common in the diets of larger individuals, but are likely ingested incidentally while consuming prey along the bottom substrate.16 Diptera larvae and adults have been reported as the most common diet item in fish of all sizes.16

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. Miscellaneous

Publications of the Museum of Zoology, University of Michigan No. 119. 131 p.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Cunningham, G.R. 1995. Life history traits and reproductive ecology of the pearl

dace, Margariscus margarita (Pisces: Cyprinidae) in the Sandhills of Nebraska.

M.S. Thesis, University of Nebraska-Omaha, Omaha, Nebraska.

4. Cunningham, G.R. 2006. Pearl Dace (Margariscus margarita): a technical

conservation assessment. USDA Forest Service, Rocky Mountain Region.

https://www.fs.usda.gov/Internet/FSE_DOCUMENTS/stelprdb5206789.pdf

5. Cunningham, G.R., R.D. Olson, and S.M. Hickey. 1995. Fish surveys of the

streams and rivers of South-central South Dakota west of the Missouri River.

Proceedings of the South Dakota Academy of Science. 74:55-64.

6. Cunninham, G. and R. Olson. 1994. Fish species collected in streams in West

River South Dakota-1994. Unpublished report to South Dakota Game, Fish and

Parks. Pierre, South Dakota 10 pp.

7. Fava, J.A, and C. Tsai. 1974. The life history of the pearl dace, Semotilus

margarita, in Maryland. Chesapeake Science 15:159-162. 261

8. Felts, E.A., and K.N. Bertrand. 2014. Conservation status of five headwater

stream specialists in South Dakota. The American Midland Naturalist 172:131-

159.

9. Langlois, T.H. 1929. Breeding habits of the northern dace. Ecology 10:161-163.

10. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

11. Lindsey, C.C. 1956. Distribution and taxonomy of fishes in the McKenzie

drainage of British Columbia. Journal of the Fisheries Research Board of Canada

13:759-886.

12. McPhail, J.D., and C.C. Lindsey. 1970. Freshwater fishes of Northwestern

Canada and Alaska. Fisheries Research Board of Canada, Ottawa. 381p.

13. Menzel, B.W. 1981. Iowa’s waters and fishes: A century and a half of change.

Proceedings of the Iowa Academy of Science 88:17-23.

14. Menzel, B.W., and M.S. Boyce. 1973. First record of the pearl dace, Semotilus

margarita (Cope), from Iowa. Iowa State Journal of Research 47:245-248.

15. Scott, W.B., and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Ottawa, Ontario, Canada. 966p.

16. Stasiak, R.H. 1978. Food, age and growth of the Pearl Dace, Semotilus margarita,

in Nebraska. The American Midland Naturalist 100:463-466.

17. Tallman, R.F., and J.H. Gee. 1982. Intraspecific resource partitioning in a

headwaters stream fish, the pearl dace, Semotilus margarita (Cope).

Environmental Biology of Fishes 7:243-249. 262

18. Whittier, T.R., and R.M. Hughes. 1998. Evaluation of fish species tolerances to

environmental stressors in lakes in the northeastern United States. North

American Journal of Fisheries Management 18:236-252.

263

Hornyhead Chub, Nocomis biguttatus (Kirtland, 1841)

Etymology and Synonyms: Nocomis = an Indian name applied by Charles Girard, a

French ichthyologist, to reference a group of fishes; biguttatus = “two spotted”.

Description: Body fusiform, robust, cylindrical anteriorly and laterally compressed posteriorly. Dorsally dark brown to olive; laterally bronze, olive to silver with dark, diffuse lateral stripe ending in a spot at the base of the caudal fin; ventrally silver to white; fins lightly pigmented. Head broad. Snout bluntly pointed. Eye large, placed laterally on upper portion of head. Mouth large, terminal; upper jaw extends to anterior edge of the eye. Barbels present, small, inconspicuous; one on each corner of the mouth.

Frenum absent. Lips fleshy. Pharyngeal tooth pattern 1,4-4,1. Gill rakers short, 9. Dorsal fin with 8 rays, nearly straight to slightly convex distal end. Adipose fin absent. Caudal peduncle moderately thick. Caudal fin moderately forked. Anal fin with 7 rays. Pelvic fins abdominal with 8 rays, insertions directly under or slightly anterior to insertion of dorsal fin. Pectoral fins with 14-17 rays. Lateral line complete with 38-45 cycloid scales in series. Spawning males develop a distinct red-orange spot posterior of the eye laterally on the head, as well as large tubercles on the dorsal side of the head and smaller tubercles on the pectoral fin rays. Juveniles similar in appearance to adults with a more pronounced lateral stripe and caudal spot.

Similar Species: Closely resembles the Creek Chub. Creek Chub have an upper jaw that extends past the anterior edge of the eye, a black spot on the anterior base of the dorsal fin present, an anal fin with 8 rays, and the pelvic fin insertions are anterior to the dorsal fin insertion. Lake Chub have a relatively small mouth and an anal fin with 8 rays. 264

Distribution and Habitat: Widespread native range in the north central glacial regions of the United States across the Missouri, upper Mississippi, northern Ohio and St.

Lawrence-Great Lakes drainages.8 Distribution of the species has decreased dramatically throughout its historical native range likely due to anthropogenic factors such as increased agricultural activity, impoundments and intermittency of stream flows, siltation and drought.8,10 Widespread in the upper Red River of the North in North Dakota, and occurs within the Big Sioux and Minnesota River basins in eastern South Dakota.1,8

Inhabit the headwaters of creeks and small rivers of low to moderate gradients with clear water, clean gravel, rubble, or sand substrates, and some aquatic vegetation which can provide refuge from large piscivores.4,8,10,14 Abundance of the species has shown to be negatively related to high gradient waters with faster velocities such as the mainstems of large rivers.5,10 Mean critical thermal maximum is 35.6 °C (96.08 °F), and mean critical oxygen concentration is reported as 1.06 mg/L.13 Juveniles are frequently encountered within algal and macrophyte beds in water with little to no current.7

Reproduction: Spawning takes place late May through early June, or when water temperatures range 16-26 °C (60.8-78.8 °F).11,14 Sexual maturity occurs at age 2-3, with males maturing at a greater length than females.7 Spawning behavior may be postponed or negatively affected by flow mediations, which can reduce the number of individuals spawning annually.10 Males put in a significant amount of time and energy into constructing nets within open areas over gravel substrate in significantly deeper and faster waters than the average stream depth and current velocities.14,15 Larger stones and pieces of gravel are carried by males in their mouth, or pushed with the dorsal surface of their heads to create a depression roughly 5-10 cm (1.97-3.94 in) deep.14 Stones and 265 gravel are continuously placed into the depression until a mount is created.14 On top of the mound, males will press and vibrate their bodies against the substrate and use their ventral side and anal fins to further form a central “cup” depression so that the nest is

“doughnut” shaped.14 Nests are often within close proximity of other nests and average

6.4 cm (0.21 ft) high, 50 cm (1.64 ft) wide and 40 cm (1.31 ft) long.14 Spawning behavior typically consists of a single female approaching a single male, the pair aligning their bodies parallel, and the males caudal peduncle wrapped around the females.14 Males flex their body and apply pressure to the females abdomen, pushing them into the “cup” area of the nest to initiate the release of eggs and milt.14 Once eggs are deposited, males cover them with more stones and pebbles to help incubate them.14 One male may occasionally spawn with multiple females.14 Fecundity of four females ranging 80-89 mm (3.15-3.50 in) SL is reported as 460-725 eggs.7 Eggs adhesive.2

Age and Growth: Mean lengths-at-age of males from Wisconsin are reported as: age-0,

35.6 mm (1.40 in) TL; age-1, 65.5 mm (2.58 in) TL; age-2, 97.9 mm (3.85 in) TL; age-3,

125.7 mm (4.95 in) TL.2 Males grow more rapidly than females.7 Capable of reaching

203.2 mm (8.00 in) TL.6 Longevity 4 years.7

Food and Feeding: Visual omnivore.3,10 Adults forage on algae and other plant material, aquatic insects, detritus and even other species of small fish.11,12 Adults are also known to consume worms, crayfish and snails.12 Juvenile diets consist of much smaller prey items including algae, zooplankton, diatoms, and small aquatic insect larvae.9,12

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor. 266

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Davis, B.J., and R.J. Miller. 1967. Brain patterns in minnows of the genus

Hybopsis in relation to feeding habits and habitat. Copeia 1967(1)1-39.

4. Hayer, C.A., S.S. Wall, and C.R. Berry, Jr. 2008. Evaluation of predicted fish

distribution models for rare fish species in South Dakota. North American Journal

of Fisheries Management 28:1259-1269.

5. Hickerson, B.T. 2018. Conservation and recovery of hornyhead chub. M.S.

Thesis, University of Wyoming, Laramie, Wyoming.

6. Hubbs, C.L., and G.P. Cooper. 1936. Minnows of Michigan. Bulletin of the

Cranbrook Institute of Science 8:1-95.

7. Lachner, E.A. 1952. Studies of the biology of the cyprinid fishes of the chub

genus Nocomis of northeastern United States. American Midland Naturalist

48:433-466.

8. Lachner, E.A., and R.E. Jenkins. 1971. Systematics, distribution, and evolution of

the Nocomis biguttatus species group (Family Cyprinidae: Pisces), with a

description of a new species from the Ozark Upland. Smithsonian Contributions

to Zoology 91:1-28.

9. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

10. Mammoliti, C.S. 2002. The effects of small watershed impoundments on native

stream fishes: a focus on the Topeka shiner and hornyhead chub. Transactions of

the Kansas Academy of Science 105:219-231. 267

11. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

12. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish.

Research Board of Canada, Bulletin 184. 966p.

13. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

14. Vives, S.P. 1990. Nesting ecology and behavior of hornyhead chub Nocomis

biguttatus, a keystone species in Allequash Creek, Wisconsin. American Midland

Naturalist 124:46-56.

15. Wisenden, B.D., A. Unruh, A. Morantes, S. Bury, B. Curry, R. Driscoll, M.

Hussein and S. Markegard. 2009. Functional constraints on nest characteristics of

pebble mounds of breeding male hornyhead chub Nocomis biguttatus. Journal of

Fish Biology 75:1577-1585.

268

Golden Shiner, Notemigonus crysoleucas (Mitchill, 1814)

Etymology and Synonyms: Notemigonus = angled back, in reference to body shape; cryso = gold, leucas = white; in reference to body coloring.

Description: Body deep, fusiform, laterally compressed with a fairly arched back.

Dorsally iridescent gold to olive; laterally silver to light olive; ventrally silver-yellow; may have an entirely silver body; fins faint yellow-orange to clear in color. Head small, flattened, triangular. Mouth small, nearly vertical, oblique; maxilla extends to nostril; snout pointed. Eye large. Barbels absent. Frenum absent. Pharyngeal tooth pattern 0,5-

5,0. Gill rakers long; 16-21. Dorsal fin with 7-9 rays, no spines; slightly falcate; insertion distinctly posterior to pelvic fin. Adipose fin absent. Caudal peduncle short, narrow.

Caudal fin deeply forked. Anal fin long, concave with 11-14 rays, no spines. Pelvic fin abdominal. Pectoral fin rays 15-18. Keel scale-less from pelvic to anal fin. Lateral line complete, strongly decurved; 42-54 cycloid scales. Peritoneum silvery with dark specks.

Intestine short; single S-shaped loop. Juveniles with dusky lateral stripe as wide as eye.

Breeding males with orange coloring on caudal, anal, and pelvic fins; small tubercles present on head and lateral sides.

Similar Species: Rudd closely resemble Golden Shiner. Rudd have 36-45 lateral line scales, red fins on adults, scales present on the ventral keel from the pelvic to anal fin, 9-

11 dorsal fin rays, and fewer gill rakers (10-13). Red Shiner have a more blunt snout, and a slightly decurved lateral line with 32-36 diamond shaped scales. Common Shiner have a slightly decurved lateral line with 36-44 diamond shaped scales with a height 3 times the width. Common Shiner also have a dorsal fin insertion roughly even with the insertion of the pelvic fins. 269

Distribution and Habitat: Widely distributed across the United States due to popularity as a bait and forage fish. Native to southeastern Canada and the eastern and central

United States.10,11 Native to drainages east and west of the Missouri River in both North

Dakota and South Dakota.8 In the Dakotas, they are abundant in lakes and ponds where they serve as a food source for larger predators such as Largemouth Bass. Commonly found throughout their range in lakes, reservoirs, ponds, creeks, and small to medium sized rivers with little flow, away from areas with fast currents. Not common in intermittent streams or larger rivers with higher flow levels.4 Prefer clear water in areas with abundant, submerged aquatic vegetation over mud, sand, or gravel substrates.

Commonly found in schools just below the surface or in the mid-water column. Able to tolerate a wide range of water temperatures with a thermal maximum of 26-40°C (78.8-

104°F).11 Also known to be tolerant to turbidity as well as relatively low dissolved oxygen concentrations.11,12,13

Reproduction: Fractional spawners, allowing for rapid repopulation after winter kill events in the northern ranges. Spawning occurs during the late spring and summer in water temperatures of 13-30°C (55-86°F), typically in the morning over beds of aquatic vegetation.3,11 Sexual maturity reached between ages 1 and 2, although can mature sooner in the south.5,11 One or two males known to court the female to the spawning grounds, often nudging with their snouts.6 Broadcast spawners. Eggs small, roughly 1-1.4 mm

(0.04-0.06 in) in diameter, and adhere to aquatic vegetation, filamentous algae, or substrates. No nest is prepared, and no parental care is given. Sometimes known to deposit eggs within nests of other fish species, including sunfish and bass species, which 270 inadvertently provide protection to the eggs.9 Fecundity known to range from 20,602-

26,079 eggs/female.3,11 Depending on water temperature, eggs hatch within 2-5 days.11

Age and Growth: Newly hatched larvae known to be 4.0-4.3 mm (0.16-0.17 in) TL.1,11

Early growth rate rapid with favorable water temperatures. Adults average 203-305 mm

(8-12 in) TL; have been known to reach 356 mm (14 in) TL. Males known to grow slower and be smaller than females.2 Adult growth rate highly dependent on water temperature and length of growing season; often grows faster in warmer climates.

Longevity roughly 8 years.

Food and Feeding: Omnivorous. Often feed just below the surface or within the mid- water column. Exhibits diel movement. Generally feed near shore and within vegetated areas during the daylight hours on filamentous algae, small terrestrial and aquatic insects and larvae, small crustaceans (cladocerans, copepods, and ostracods). In the evening, they filter feed with the help of their long gill rakers, taking place in open and deeper water, primarily on zooplankton.7 Small snails and small fish are sometimes consumed by larger individuals.

Literature Cited:

1. Buynak, G.L. and H.W. Mohr Jr. 1980. Larval development of golden shiner and

comely shiner from northeastern Pennsylvania. Progressive Fish-Culturist 42:206-

211.

2. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

3. Clemment, T. and N. Stone. 2010. Golden shiner egg production during a

spawning season. North American Journal of Aquaculture 72:272-277. 271

4. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

5. Dobie, J.R., O.L. Meehean, and G.N. Washburn. 1948. Propagation of minnows

and other bait species. U.S. Fish and Wildlife Service Circular 12, Washington,

District of Columbia, USA.

6. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

7. Hall, D.J., E.E. Werner, J.F. Gilliam, G.G. Mittelbach, D. Howard, C.G. Doner,

J.A. Dickerman, and A.J. Stewart. 1979. Diel foraging behavior and prey

selection in the golden shiner (Notemigonus crysoleucas). Journal of the Fisheries

Research Board of Canada 36:1029-1039.

8. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry Jr. 2007.

Zoogeographic patterns and faunal change of South Dakota fishes. Western North

American Naturalist 67(2):161-184.

9. Kramer, R.H., and L.L. Smith. 1960. Utilization of nests of largemouth bass,

Micropterus salmoides, by golden shiners, Notemigonus crysoleucas. Copeia

1960: 73-74.

10. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. Publication 1980-

12, North Carolina Biological Survey, North Carolina State Museum of Natural

History, Raleigh, North Carolina, USA. 272

11. Stone, N.M., A.M. Kelly, and L.A. Roy. 2016. A fish of weedy waters: golden

shiner biology and culture. Journal of the World Aquaculture Society, 47(2):152-

200.

12. Trebitz, A.S., J.C. Brazner, V.J. Brady, R. Axler, and D.K. Tanner. 2007.

Turbidity tolerances of Great Lakes coastal wetland fishes. North American

Journal of Fisheries Management 27:619-633.

13. Whittier, T.R. and R.M. Hughes. 1998. Evaluation of fish species tolerances to

environmental stressors in lakes in the northeastern United States. North

American Journal of Fisheries Management 18:236-252.

273

Pugnose Shiner, Notropis anogenus (Forbes, 1885)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; anogenus = “without chin”.

Description: Body fusiform, slender, slightly laterally compressed. Dorsally dusky olive, tan, gold or bronze with scales outlined in slightly darker pigment creating a

“crosshatched” pattern; laterally silver with prominent black lateral stripe present extending from the snout, through the eye to a wedge shaped spot at the base of the caudal fin; adults without concentrated pigment on some lateral scales causing some scales to appear missing; ventrally silver to white; fins generally clear to lightly pigmented. Head moderately large. Snout blunt. Eye moderately large, greater than the length of snout, placed laterally on head. Mouth small, superior and oblique, nearly vertical; upper jaw extends posteriorly below the nostril. Barbels absent. Frenum absent.

Pharyngeal tooth pattern 0,4-4,0. Gill rakers short, 8. Dorsal fin with 8-9 rays and straight distal end. Adipose fin absent. Caudal peduncle slightly elongate, uniform in thickness.

Caudal fin moderately forked. Anal fin with 7-8 (may sometimes be up to 10) rays and straight distal end. Pelvic fins abdominal with 8 rays; insertions slightly anterior to insertion of dorsal fin. Pectoral fins with 12-14 rays. Lateral line complete with 34-37 cycloid scales in series; scales on the anterior lateral side of body as deep as they are wide. Spawning males develop small tubercles on the dorsal side of head, as well as on the pectoral and pelvic fin rays. Sexually dimorphic species; pelvic fin of males extends to or slightly beyond cloaca; females pelvic fin does not reach the cloaca. Juveniles similar to adults. 274

Similar Species: Easily distinguished from other species of Notropis by the small, superior, oblique, and nearly vertical mouth. May be confused with the Blacknose Shiner,

Brassy Minnow and Fathead Minnow. Blacknose Shiner display dark borders on the lateral stripe pores that expand into crescent-shaped vertical bars anteriorly, a snout that slightly extends beyond the mouth, a lower jaw that extends to the anterior edge of the eye, and an incomplete lateral line. Brassy Minnow have a snout that slightly protrudes past the mouth, and pelvic fin insertions that are slightly posterior to the dorsal fin insertion. Fathead Minnow have a dorsal fin with a rounded distal end, pectoral fins with

15-16 rays, and an incomplete lateral line with 40-54 cycloid scales in series.

Distribution and Habitat: Considered one of the rarest cyprinids in the northern United

States, Pugnose Shiner are characterized by small and relatively isolated populations.1,8,9

Native range is fragmented across the Upper Mississippi River, Red River of the North, and Great Lakes basins, from western New York and eastern Ontario to southeastern

North Dakota.3,5 Reported from the Sheyenne and Turtle Rivers within the Red River basin in North Dakota.9 Not reported in South Dakota. Pugnose Shiner appear to be declining throughout their native distribution due to several factors such as their extreme sensitivity to turbidity, shoreline erosion and development which cause reduced amounts of aquatic vegetation, and the introduction of invasive species.1,3,4,7,10,11 An abundance of submerged aquatic vegetation is an important variable to the species life history as it provides cover from predators, foraging opportunities, and likely acts as a reproductive site.2,3 Preferred habitat includes clear, quiet, glacial lakes and stagnant channels of low gradient streams with abundant submerged vegetation and clean sand or organic debris substrate.3,11 275

Reproduction: Information on spawning and reproductive behavior of the Pugnose

Shiner is limited. Spawning in Wisconsin is recorded to take place from mid-May through July, or when water temperatures reach 21-29 °C (69.8-84.2 °F).2 Lithophilic spawner.3 No nest is constructed, and no parental care is given. Gravid females are capable of producing a total of 530-1,275 eggs, and likely release their eggs more than once during a spawning season.2 Eggs yellow and roughly 0.5-1.3 mm (0.02-0.05 in) in diameter.2

Age and Growth: Mean length of age-0 individuals from Ontario, Canada reported as

24.1 mm (0.95 in) TL.3,6 Mean lengths-at-age of individuals from Wisconsin are reported as: age-1, 42 mm (1.65 in) TL; age-2, 46.3 mm (1.82 in) TL; age-3, 52.5 mm (2.07 in)

TL.2 Females capable of reaching 60 mm (2.36 in) TL, and males capable of reaching 50 mm (1.97 in) TL.3 Longevity 3 years.3

Food and Feeding: Little information is known about the diet and feeding habits of the

Pugnose Shiner. Diets of Pugnose Shiner from Ontario, Canada consist of small, minute plant material such as Chara and filamentous green algae, cladocerans such as Daphnia,

Bosmina, and Chydorus, caddisfly larvae, and small leeches.2,3 Foraging mainly takes place within the midwater.2

Literature Cited:

1. Bailey, R.M. 1959. Distribution of the American cyprinid fish (Notropis

anogenus). Copeia (1959):119-123.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 276

3. COSEWIC. 2002. COSEWIC assessment and update status report on the pugnose

shiner Notropis anogenus in Canada. Committee on the Status of Endangered

Wildlife in Canada. Ottawa. Vi + 15pp.

4. DFO. 2010. Recovery potential assessment of pugnose shiner (Notropis

anogenus) in Canada. DFO Canadian Science Advisory Secretariat Science

Advisory Report 2010/025.

5. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

6. Leslie, J.K., and C.A. Timmins. 2002. Description of age 0 juvenile pugnose

minnow Opsopoedus emiliae (Hay) and pugnose shiner Notropis anogenus

Forbes in Ontario. Canadian Technical Report of Fisheries and Aquatic Sciences

2397.

7. Lyons, J. 1992. Using the index of biotic integrity (IBI) to measure environmental

quality in warmwater streams of Wisconsin. General Technical Report NC-149.

U.S. Department Agriculture, Forest Service. North Central Forest Experimental

Station, St. Paul MN.

8. McCusker, M.R., N.E. Mandrak, B. Egeh, and N.R. Lovejoy. 2014. Population

structure and conservation genetic assessment of the endangered pugnose shiner,

Notropis anogenus. Conservation Genetics 15:343-353.

9. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks. 277

10. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

11. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

278

Emerald Shiner, Notropis atherinoides (Rafinesque, 1818)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; atherinoides =

Greek for the words “atherina”, meaning “silverside”, and “oides”, a Greek suffix meaning “resemblance”.

Description: Body fusiform, streamlined, elongate, and laterally compressed. Dorsal side lightly pigmented, yellowish-olive to iridescent blue-green; melanophores only along distal edges of anterior dorsolateral scales (creating an outlined appearance); laterally silver with faint silvery lateral stripe continuous onto the opercle and then fading near the anterior end of the dorsal fin; ventrally silver to white; fins clear to lightly pigmented.

Head small. Snout short and bluntly rounded; length less than two-thirds the distance from the posterior margin of the eye to the posterior margin of the head. Eyes large; diameter greater than the snout length; placed laterally on head. Mouth large, oblique, and terminal. Teeth absent. Barbles absent. Frenum absent. Lips and center of chin deeply pigmented. Gill rakers 10-12. Dorsal fin pointed with 8 rays; anterior fin ray reaches past the posterior fin ray when depressed; posterior fin ray is less than one half the length of the longest ray; insertion distinctly posterior to pelvic fin insertions.

Adipose fin absent. Caudal peduncle moderately elongate, uniform in thickness. Caudal fin moderately forked. Anal fin with 10-13 rays. Pelvic fins abdominal with 8-9 rays.

Pectoral fins with 14-16 rays. Lateral line complete with 35-43 cycloid scales in series.

Spawning males and females similar to non-spawning appearance. Juveniles similar to adults.

Similar Species: Closely resembles the River Shiner, Channel Shiner, Carmine Shiner,

Silverband Shiner and Bigmouth Shiner. River Shiner have a dark, well-defined mid- 279 dorsal stripe present that surrounds the dorsal fin base, have a fewer number of lateral line scales in the series (32-36), and have an anal fin with 7 rays. Channel Shiner have scales on their dorsal side that are outlined in darker pigment creating a prominent

“crosshatched” pattern, and have an anal fin with 8 rays. Carmine Shiner are laterally silver to iridescent blue, have a faint rosy pink lateral stripe extending onto the opercle and cheek, and have a snout that is slightly greater in length than the diameter of the eye.

Silverband Shiner have a dorsal fin insertion only slightly posterior to the pelvic fin insertion. Bigmouth Shiner have a snout that overhangs the mouth, and have eyes that are directed upward with the pupils visible when viewed from up above.

Distribution and Habitat: Emerald Shiner have one of the largest ranges of all North

American minnows. In the U.S. their native range extends from the Missouri River basin in the west, to the Mississippi, Hudson, St. Lawrence-Great Lakes, and Mackenzie River drainages in the east, and south throughout the southern Mississippi and Gulf Slope drainages to the Gulf of Mexico. Widely distributed throughout the Dakotas, and is likely the most abundant minnow species in the Missouri and Mississippi rivers.11 Primarily occurs west of the Missouri River in North Dakota throughout the Little Missouri, Knife,

Heart and Cannonball River drainages, but is also known to occur in the Souris, James,

Sheyenne and Red River of the North drainages.14 Occurs throughout the vast majority of the main Missouri River tributaries and their watersheds in South Dakota, as well as the

Upper Minnesota and Red River of the North drainages.7 Often found schooling near the surface of rivers, large streams, reservoirs and lakes with clear to slightly turbid water.

The species is known to avoid areas with aquatic vegetation.3 Emerald Shiner are able to 280 tolerate lower concentrations of dissolved oxygen and higher turbidity, however they generally do not thrive in waters with higher temperatures.2,8,12

Reproduction: Spawning primarily occurs throughout late spring and early summer

(although they have been recorded to spawn in August), and is known to be temperature dependent, beginning when water temperatures reach roughly 22.2-24.0°C (72.0-

75.2°F).5,6,15 Sexual maturity begins to take place at 55-60 mm (2.2-2.4 in) TL for males, and roughly 65 mm (2.6 in) TL for females, with the majority of individuals mature by age-2.5 Spawning behavior spans a period of roughly 4-6 weeks, and consists of mature adults gathering in large schools at night near the surface over sand or gravel substrate.5

Smaller males tend to pursue larger females, and once paired up, the male presses his body against the female to appear as if they are interlocking their pectoral fins.5 The act of spawning may take place more than once during the season. Eggs are demersal, nonadhesive, and range from 0.21-0.8 mm (0.008-0.031 in) in diameter.1,4 Fecundity reports from Canada ranged 868-8733 eggs per female, with the number of eggs increasing with the length, weight, and gonad weight of the female.4 In contrast, an age-1

69 mm (2.7 in) TL female from Wisconsin was reported to produce 2,990 eggs, and an age-2 75 mm (2.95 in) TL female produced 2,040 eggs.1 Hatching takes place within 24-

32 hours depending on water temperature.1

Age and Growth: Growth rates are known to be rapid initially. In Lewis and Clark Lake,

South Dakota, average lengths from the first and second annulus formation are reported as 66 mm (2.6 in) TL and 84 mm (3.3 in) TL.6 Females seem to attain larger sizes than males within the first year of growth, and continue to be larger throughout their life.6

Maximum growth rates have been recorded at 28.9°C (84.0°F).9 Longevity age-4, 281 although majority of individuals only reach age-2, with the older individuals being female.6,13

Food and Feeding: Juveniles feed heavily on rotifers and algae.6 Adults feed primarily on larger zooplankton such as cladocerans and copepods, but are also known to feed on adult and larval insects, annelids, smaller fish, and algae.2,6 Adults have been observed feeding on drifting invertebrates during the day in the upper water column in rivers, as well as rising to the surface to consume insects.10

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

4. Campbell, J.S., and H.R. MacCrimmon. 1970. Biology of the emerald shiner,

Notropis atherinoides Raf., in Lake Simcoe, Canada. Journal of Fisheries Biology

2:259-273.

5. Flittner, G.A. 1964. Morphometry and life history of the emerald shiner Notropis

atherinoides Rafinesque. PhD dissertation. University of Michigan, Ann Arbor.

6. Fuchs, E.H. 1967. Life history of the emerald shiner, Notropis atherinoides in

Lewis and Clark Lake, South Dakota. Transaction of the American Fisheries

Society. 96: 247-256. 282

7. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry Jr. 2007.

Zoogeographic patterns and faunal change of South Dakota fishes. Western North

American Naturalist 67:161-184.

8. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

9. McCormick, J.H., and C.F. Kleiner. 1976. Growth and survival of young-of-the-year

emerald shiners (Notropis atherinoides) at different temperatures. Journal of the Fisheries

Board of Canada, 33: 839-842.

10. Mendelson, J. 1975. Feeding relationships among species of Notropis (Pisces:

Cyprinidae) in a Wisconsin stream. Ecological Monographs 45:199-232.

11. Neuman, R.M., and D.W. Willis. 1994. Guide to the Common Fishes of South

Dakota. Extension Circulars. Paper 511.

12. Pflieger, W.L. 1997. The Fishes of Missouri, revised edition. Missouri

Department of Conservation, Jefferson City.

13. Ross, S.T. 2001. The inland fishes of Mississippi. Mississippi Department of Wildlife,

Fisheries, and Parka, and University Press of Mississippi, Jackson.

14. Russel, G.W. 1975. Distribution of fishes in North Dakota drainages affected by

the Garrison Diversion Project. M.S. Thesis, University of North Dakota. 100p.

15. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966pp.

283

River Shiner, Notropis blennius (Girard, 1856)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; blennius = Greek for blennos, meaning “mucus” or “slime”.

Description: Body fusiform, robust, slightly laterally compressed. Dorsally tan to dusky silver with scales faintly outlined in darker pigment, and dark, well-defined mid-dorsal stripe present that surrounds the dorsal fin base; laterally silver with no distinct markings; ventrally silver to white; fins clear. Head moderately large. Snout bluntly rounded, slightly protruding beyond the mouth. Eye large, placed laterally on head. Mouth large, terminal, and slightly oblique; upper jaw extends to or slightly beyond anterior end of eye and is greater in length than the diameter of the eye. Barbels absent. Frenum absent.

Pharyngeal tooth pattern variable between 1,4-4,1 and 2,4-4,2. Gill rakers short, 6-9.

Dorsal fin with 8 rays and pointed tip; distal end nearly straight to slightly convex.

Adipose fin absent. Caudal peduncle slightly elongate, uniform in thickness. Caudal fin moderately forked. Anal fin with 7 rays and straight distal end. Pelvic fins abdominal with 8 rays; insertions directly inferior to dorsal fin insertion. Pectoral fins with 13-15 rays. Lateral line complete and slightly decurved with 32-36 cycloid scales in series.

Spawning males develop small tubercles on the dorsal side of the head, the pectoral fins, and on the anterior ends of the dorsal and anal fins. Juveniles similar to adults.

Similar Species: Closely resembles the Sand Shiner, Emerald Shiner, Bigmouth Shiner,

Channel Shiner and Silverband Shiner. Sand Shiner display small, paired, dark specks or

“mouse tracks” outlining the lateral line and a faint, dusky spot sometimes present at the base of the caudal fin. Emerald Shiner have an anal fin with 10-13 rays, and a dorsal fin insertion that is distinctly posterior to the insertion of the pelvic fins. Bigmouth Shiner 284 have a large, slightly subterminal mouth with the lower jaw extending to the middle of the eye, and the length of the upper jaw greater than the eye diameter. Channel Shiner have a lateral stripe expanding ventrally at the base of the caudal fin. Silverband Shiner have an anal fin with 8-9 rays, and pelvic fins with 9 rays and insertions slightly posterior to the dorsal fin insertion.

Distribution and Habitat: Native to the Hudson Bay drainage from Manitoba to

Alberta, Canada in the north, south throughout the Red River, Mississippi, Middle

Missouri, Ohio, and Arkansas River basins.5 Occurs within the Cannonball, Heart, and

Knife River watersheds west of the Missouri River in North Dakota, as well as the

Souris, Sheyenne, and Red River watersheds on the eastern end of the state.6 Confined to the lower Middle Missouri River, and the lower reaches of the Vermillion, James, and

Niobrara rivers in southeastern South Dakota.1 Typically found schooling within the midwaters of large lakes, large, broad channeled rivers, and the lower parts of main tributaries with clear to turbid water and moderate to swift current over gravel and sand bars.2,3,7,8 Capable of withstanding prolonged periods of increased turbidity, and typically avoids areas with strong current.7

Reproduction: Little information is known about the River Shiners spawning and reproductive habits. Spawning season is prolonged and takes place from June to

August.2,3,7 Males reach sexual maturity at age-1, and females at age-2.2 Spawning occurs over sand and gravel substrate.8 Fractional spawners, with females producing multiple clutches of eggs over the spawning season.4 Mature females ranging 48.3-87.9 mm (1.90-

3.46 in) SL from the Upper Mississippi River produced 436-2,754 eggs per clutch.4 285

Clutch size positively correlated with the length of the female.4 Eggs yellow in color, and roughly 0.89 mm (0.04 in) in diameter.4

Age and Growth: Mean lengths-at-age of individuals from the Mississippi River in

Wisconsin are reported as: age-0, 37.1 mm (1.46 in) TL; age-1, 58.4 mm (2.30 in) TL; age-2, 76.5 mm (3.01 in) TL; age-3, 87.3 mm (3.44 in) TL; age-4, 97.3 mm (3.83 in) TL.2

Capable of reaching 132 mm (5.20 in) long.8 Longevity 4 years.2

Food and Feeding: Omnivorous. Adult diets primarily consist of aquatic insects including larvae and adult Chironomids Trichopterans, and Corixids, but may also consume algae, microcrustacea, other plant material, and terrestrial insects.3,9

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

4. Hatch, J.T., and E.E. Elias. 2002. Ovarian cycling, clutch characteristics, and

oocyte size of the river shiner Notropis blennius (Girard) in the Upper Mississippi

River. Journal of Freshwater Ecology 17:85-92.

5. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp. 286

6. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

7. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

8. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

9. Whitaker, J.O. Jr. 1977. Seasonal changes in food habits of some cyprinid fishes

from the White River at Petersburg, Indiana. The American Midland Naturalist

97:411-418.

287

Bigmouth Shiner, Notropis dorsalis (Agassiz, 1854)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; dorsalis = Latin for

“pertaining to the back”.

Description: Body fusiform, slender, slightly laterally compressed. Dorsally brown, tan or olive with dorsal scales outlined in darker pigment creating a “crosshatched” pattern; prominent, narrow mid-dorsal stripe present and continuous on each side of dorsal fin; laterally silver with small, paired, dark specks or “mouse tracks” outlining the lateral line; ventrally silver to white; fins with little to no pigment. Head elongate with lower surface distinctly flattened. Snout bluntly pointed; slightly overhangs the mouth. Eye large, placed laterally on head, but directed upwards with lower margins of pupils visible when fish is viewed from above. Mouth large, slightly subterminal with lower jaw extending to the middle of the eye; length of upper jaw greater than diameter of the eye. Barbels absent. Frenum absent. Pharyngeal teeth 1,4-4,1. Gill rakers short, roughly 8. Dorsal fin with 8 rays. Adipose fin absent. Caudal peduncle slender, elongate. Caudal fin moderately forked. Anal fin with 8 rays and straight distal end. Pelvic fins abdominal with 8 rays, insertions directly under or slightly anterior to insertion of dorsal fin.

Pectoral fins with 14-15 rays. Lateral line complete with 32-39 cycloid scales in series.

Spawning males develop small tubercles on the dorsal and lateral sides of the head, as well as on the pectoral fin rays. Juveniles similar to adults.

Similar Species: Closely resembles the Sand Shiner, Emerald Shiner and Spottail Shiner.

Sand Shiner have an anal fin with 7 rays, an upper jaw that is shorter than the diameter of the eye, and eyes not directed upward. Emerald Shiner have an anal fin with 10-13 rays, and a dorsal fin insertion that is distinctly posterior to the insertion of the pelvic fins. 288

Spottail Shiner have a prominent, dark, black spot present at base of caudal fin that is as large or larger than the size of the pupil.

Distribution and Habitat: Native in the west throughout the Platte River basin in eastern Wyoming and northeastern Colorado, to eastern North Dakota and South Dakota in the north, and east throughout the Upper Mississippi, St. Lawrence-Great Lakes and

Hudson Bay drainages to Illinois and Missouri with native disjunct populations present in

New York and West Virginia.3 Inhabits the James, Sheyenne, Souris, Cannonball, Heart and Little Missouri Rivers in North Dakota, as well as the Minnesota tributaries of the

Red River.6 Widely distributed and occurs throughout all the main river drainages of eastern South Dakota, as well as the Niobrara, White, Bad and Grand River drainages west of the Missouri River.1 Often found over sand bars or small, shallow pooled areas free of vegetation over sand and silt substrate in small creeks or open prairie-like streams of moderate gradient with clear to turbid water, slight current, and permanent flow.2,3,4,7,9

Also inhabits sloughs and lakes.2 Often found schooling with other minnow species such as the Sand Shiner.4,7 Abundance tends to decrease when stream width reaches >3.05 m

(10 ft).9 At a water temperature of 26 °C (78.8 °F), mean critical dissolved oxygen concentration is reported as 1.02 mg/L.8 Mean thermal maximum 36.6 °C (97.9 °F).8

Reproduction: Little information is known about the Bigmouth Shiners reproductive habits, as the act of spawning has never been observed. It is suggested that the spawning habits are similar to other shiner species, in which spawning takes place midstream over sand substrate with eggs developing as they drift downstream.2,5 Sexual maturity is reached at age-1 in Missouri, and at age-2 in Wisconsin.7,2 In the northern part of the species range, spawning begins in May, and likely extends to August.2 In the southern 289 part of its range, spawning has been observed from June to July.7 Fecundity of a 61 mm

(2.40 in) TL age-2 female from Wisconsin reported as 1,000 eggs, and approximately

1,275 eggs from an age-2, 66 mm (2.60 in) TL female.2 Eggs roughly 0.9-1.0 mm (0.39-

0.04 in) in diameter and yellow in color.2

Age and Growth: Larvae from Missouri are roughly 20-41 mm (0.8-1.6 in) at age-0.7

Mean lengths-at-age of Bigmouth Shiners from Wisconsin are reported as: age-0, 44.8 mm (1.76 in) TL; age-1, 60.9 mm (2.40 in) TL; age-2, 67.2 mm (2.65 in) TL.2 Capable of reaching 80 mm (3.15 in) TL.2 Longevity 4 years.3

Food and Feeding: Benthic omnivores, forging on small aquatic insects such as

Ephemeropterans and Dipterans, as well as plant material, detritus, and zooplankton.2,7,10

Sight is presumed to be less important than taste when foraging.7 In an aquarium setting, individuals have been observed rapidly swimming along the bottom taking in mouthfuls of sand, sorting out the food particles in the mouth, and discarding the undesired particles through the mouth or gill openings.7

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. Miscellaneous

Publications of the Museum of Zoology, University of Michigan No. 119. 131 p.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

4. Mendelson, J. 1975. Feeding relationships among species of Notropis (Pisces:

Cyprinidae) in a Wisconsin stream. Ecological Monographs 45:199-230. 290

5. Moore, G.A. 1944. Notes on the early life history of Notropis girardi. Copeia

1944: 209-214.

6. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

7. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

8. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

9. Starrett, W.C. 1950a. Distribution of the fishes of Boone County, Iowa with

special reference to the minnows and darters. The American Midland Naturalist

43:112-127.

10. Starrett, W.C. 1950b. Food relationships of the minnows of the Des Moines

River, Iowa. Ecology 31:216-233.

291

Blacknose Shiner, Notropis heterolepis (Eigenmann & Eigenmann, 1893)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; heterolepis = Greek for “various scale”, referring to the varying shape of the scales.

Description: Body fusiform, slender. Dorsally tan to olive with dorsal scales outlined in darker pigment creating a “crosshatched” pattern; laterally olive to bronze with prominent, dark, dusky lateral stripe extending onto snout; dark borders of lateral stripe pores expanded into crescent-shaped vertical bars anteriorly; scales directly above lateral stripe lighter in color; ventrally silver to white; fins with little to no pigment. Head moderate. Snout bluntly rounded, slightly extending beyond the mouth. Eye moderately large, placed laterally on head. Mouth small, terminal to slightly subterminal and slightly oblique with lower jaw extending to anterior edge of eye. Barbels absent. Frenum absent.

Pharyngeal teeth 0,4-4,0. Dorsal fin with 8 rays. Adipose fin absent. Caudal peduncle slender, slightly elongate. Caudal fin forked. Anal fin with 7-8 rays and straight distal end. Pelvic fins abdominal with 8 rays, insertions directly under insertion of dorsal fin.

Lateral line incomplete with 33-37 cycloid scales in series. Spawning males with small tubercles present on dorsal side of head. Juveniles similar to adults.

Similar Species: Resembles the Brassy Minnow, Topeka Shiner and Pugnose Shiner.

Brassy Minnow display a faint, dark mid-dorsal stripe, lack any prominent

“crosshatched” patterning on the dorsal side, lack any dark borders on lateral stripe pores that expand into crescent-shaped vertical bars anteriorly, and have pelvic fin insertions that are slightly posterior to the dorsal fin insertion. Topeka Shiner have a deeper, laterally compressed body with small, paired dark specks or “mouse tracks” outlining the lateral line, and have a triangular/ chevron shaped black spot present at the base of the 292 caudal fin. Pugnose Shiner have scales on the anterior lateral side of the body that are as deep as they are wide, and a lateral stripe without pigment on some of the scales giving a

“zig-zag” appearance.

Distribution and Habitat: Native range broadly distributed over the Atlantic, Hudson

Bay, Great Lakes and Mississippi River basins.10 The species has been extirpated from much of its southern historical distribution, especially in the Midwest due to intensive row-crop agriculture and other anthropogenic disturbances including wetland loss, increased turbidity, and siltation from pollution and erosion.2,6,7,12 Blacknose Shiner are extremely rare and may soon be extirpated from South Dakota as they are limited to the perennial and spring-fed streams of the Keya Paha River basin and have disjunct or peripheral populations in the Big Sioux, James, Minnesota, Sheyenne and Red River basins.3,4,6,9 Frequently school in the midwaters of clear to moderately clear glacial lakes, and small, quiet, prairie streams with pool and run sequences.10 Often associated with considerable amounts of aquatic vegetation and organic debris, sand, gravel or rock substrate.10,11,12 Vegetation is likely important for nursery and foraging habitat.12 Likely intolerant of prolonged periods of high turbidity and siltation.11

Reproduction: Little information is known about the reproductive and spawning habits of the Blacknose Shiner. Sexual maturity generally occurs when individuals reach >33 mm (1.30 in) SL.12 Spawning in Missouri reportedly takes place June through July.11

Spawning in Illinois lakes occurs from late April or early May through late June, with females remaining in reproductive condition for roughly 2-4 weeks.12 This extended spawning period indicates that females are multiple clutch spawners.12 Clutch size of mature females ranges 98-330 eggs, with a mean of 167 eggs.12 Clutch size is known to 293 be positively related to the size of the female, however clutch size decreases over time indicating that larger clutches are laid earlier in the spawning season, especially for larger females.12 It is likely that no parental care is given.12 Eggs yellow in color, and range

0.80-0.88 mm (0.031-0.035 in) in diameter.1,12

Age and Growth: Mean lengths-at-age of individuals from Wisconsin are reported as: age-0, 30.6 mm (1.20 in) TL; age-1, 48.8 mm (1.92 in) TL.1 Mean lengths-at-age of individuals collected in October from Michigan are reported as: age-0, 35.9 mm (1.41 in)

SL; age-1, 46.1 mm (1.82 in) SL.5 Capable of reaching 81 mm (3.19 in) SL.8,11 Longevity

2 years, with many individuals likely dying before or during their second winter of life.1,5

Food and Feeding: Little information is known about the feeding habits of the

Blacknose Shiner. A subterminal mouth suggests the species is primarily a benthic feeder.1 A group of individuals in Illinois lakes opportunistically foraged during the mornings in spring, and throughout the morning and night during the summer.12 Total consumption was generally lower in the summer and fall months compared to spring.12

Diets primarily consist of Chydorid and Bosminid Cladocerans, Ostracods, Copepods,

Ephemeropterans, Dipterans, plant material, arachnids and possibly fish eggs.1,12

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Cross, F.B., and R.E. Moss. 1987. Historic changes in fish communities and

aquatic habitat in plains streams of Kansas, p. 155-165 in W.J. Matthews and

D.C. Heins (editors). Community and evolutionary ecology of North American

stream fishes. University of Oklahoma Press, Norman. 310p. 294

3. Cunningham, G. and R. Olson. 1994. Fish species collected in streams in West

River South Dakota-1994. Unpublished report to South Dakota Game, Fish and

Parks. Pierre, South Dakota 10 pp.

4. Cunningham, G.R., R.D. Olson, and S.M. Hickey. 1995. Fish surveys of the

streams and rivers of South-Central South Dakota west of the Missouri River.

Proceedings of the South Dakota Academy of Science. 74:55-64.

5. Emery, L., and D.C. Wallace. 1974. The age and growth of the blacknose shiner,

Notropis heterolepis Eigenmann and Eigenmann. The American Midland

Naturalist 91:242-243.

6. Felts, E.A., and K.N. Bertrand. 2014. Conservation status of five headwater

stream specialists in South Dakota. The American Midland Naturalist 172:131-

159.

7. Hoagstrom, C.W., C.A. Hayer, J.G. Kral, S.S. Wall, and C.R. Berry. 2006. Rare

and declining fishes of South Dakota: a river drainage scale perspective.

Proceedings of the South Dakota Academy of Science 85:171-211.

8. Hubbs, C.L., and K.F. Lagler. 1949. Fishes of Isle Royale, Lake Superior,

Michigan. University of Michigan Academy of Science 33(1947):73-133.

9. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

10. Page, L.M., and B.M. Burr. 1991. A field guide to freshwater fishes of North

America north of Mexico. Houghton Mifflin Company, Boston, Massachusetts.

432p. 295

11. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

12. Roberts, M.E., B.M. Burr, and M.R. Whiles. 2006. Reproductive ecology and

food habits of the blacknose shiner, Notropis heterolepis, in northern Illinois. The

American Midland Naturalist 155:70-83.

296

Spottail Shiner, Notropis hudsonius (Clinton, 1824)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; hudsonius = named in reference to the Hudson River.

Description: Body fusiform, slightly robust, laterally compressed posteriorly. Dorsally olive, tan, or dusky silver; laterally silver with no distinct markings; ventrally silver to white; fins generally clear; prominent, dark, black spot present at base of caudal fin (may be diffuse or absent in larger adults from turbid waters); as large or larger than size of pupil. Head small. Snout bluntly rounded, slightly protruding beyond mouth. Eye large, placed laterally on head; diameter greater than length of upper jaw. Mouth small, terminal to slightly subterminal with upper jaw extending to anterior edge of eye. Barbels absent. Frenum absent. Pharyngeal tooth pattern variable between 2,4-4,2 and 0,4-4,0.

Gill rakers short, 9. Dorsal fin with 8 rays and pointed tip. Adipose fin absent. Caudal peduncle moderately short, slender. Caudal fin forked. Anal fin with 8 rays and slightly concave distal end. Pelvic fins abdominal with 8 rays; insertions directly inferior to insertion of dorsal fin. Pectoral fins with 13-15 rays. Lateral line complete with 36-42 cycloid scales in series. Spawning males develop minute tubercles on dorsal and lateral sides of head, dorsal side of body, as well as anterior rays of the pectoral fins. Spawning females develop a less numerous amount of tubercles on the dorsal sides of the head and body. Juveniles similar to adults, sometimes with basiocaudal spot more distinct.

Similar Species: Closely resembles the Emerald Shiner, Sand Shiner, Channel Shiner, and River Shiner. Easily distinguished from these species by the presence of a dark, black spot present at the base of the caudal fin. 297

Distribution and Habitat: Widely distributed native distribution throughout North

America from the Mackenzie River system in the Northwest Territories of Canada in the northwest, and southeast through the Hudson Bay drainage, the upper eastern Mississippi

River drainage, the St. Lawrence-Great Lakes basin to the Atlantic Slope drainages.6

Present in the Cannonball, Heart, Missouri, Souris, James, Sheyenne and Red River basins of North Dakota.7 Occurs within the Cheyenne, Belle Fourche, Grand, Missouri,

James, Big Sioux and Minnesota River drainages in South Dakota.1 Spottail Shiners were introduced and stocked into Lake Oahe in 1973 as means of a forage fish.2 Inhabits the shoal waters of large lakes, glacial lakes, rivers and large creeks in areas with clear water, little to moderate current, little to moderate amounts of aquatic vegetation, and sand or gravel substrate.3,10 Prefers the shallow waters of lakes, however have been collected at depths up to 31m (101.7 ft).11,12 In Lake Oahe, Spottail Shiners appear to make diel on/offshore movements.9

Reproduction: Prolific spawners.9 Spawning takes place late May through early June, but may continue into August.5 Age at sexual maturity is a function of growth rate, with most individuals reaching maturity once they have reached roughly 68 mm (2.68 in)

TL.8,12 Males reach sexual maturity at slightly smaller lengths than females.12 Many individuals migrate to shallower depths prior to spawning.12 Fecundity generally increases with increasing length of the female.12 Ten females from Lake Michigan ranging 97-131 mm (3.82-5.16 in) TL contained 915-3,709 eggs, however females are capable of producing up to 8,898 eggs.12 Eggs yellow in color.12

Age and Growth: Females exhibit faster growth rates and generally attain greater lengths than males.12 In Lake Oahe, up to 40% of the species total growth is achieved 298 between ages 0 and 1.9 Mean lengths-at-age of individuals from Lake Oahe in July are reported as: age-0, 36.3 mm (1.43 in) TL; age-1, 81.2 mm (3.20 in) TL; age-2, 107.0 mm

(4.21 in) TL; age-3, 122 mm (4.80 in) TL.9 Mean lengths-at-age of females from southeastern Lake Michigan are reported as: age-1, 63 mm (2.48 in) TL; age-2, 97 mm

(3.82 in) TL; age-3, 114 mm (4.49 in) TL; age-4, 123 mm (4.84 in) TL; age-5, 131 mm

(5.16 in) TL.12 Individuals occupying deeper depths were seen to be older and greater in length than individuals from shallower depths.12 Capable of reaching 143 mm (5.63 in)

TL.11 Longevity 5 years.12

Food and Feeding: Opportunistic omnivores.5,9 In Lake Oahe, the most commonly consumed prey items were Daphnia, chironomid larvae, and adult Diptera.9 A shift from feeding on zooplankton to insects occurred when abundance of zooplankton was low.9 In

Lake Michigan, Spottail Shiners primarily consumed immature midges such as

Chironomids, but also foraged on Pontoporeia, zooplankton (mainly Copepods and

Cladocerans), fingernail clams, and fish eggs from Alewives and Trout-perch.11 Feeding on midges mainly took place at shallower depths, and feeding on Pontoporeia took place in deeper depths.11 Individuals have also been known to consume snails, leeches, ostracods, terrestrial insects and caddisfly larvae but in much smaller quantities.11 In Lake

Manitoba, fry consumed small plankton and Diptera larvae.4

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Barnes, M.E. 2007. Fish hatcheries and stocking practices: past and present. Pages

267-294 in C. Berry, K. Higgins, D. Willis, and S. Chipps, editors. History of 299

fisheries and fishing in South Dakota. South Dakota Department of Game, Fish

and Parks, Pierre.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Bernard, D.J. 1972. Ecological divergence between emerald and spottail shiners

(Notropis) in Lake Manitoba. M.S. Thesis, University of Manitoba, Winnipeg,

Manitoba, Canada.

5. Griswold, B.L. 1963. Food and growth of spottail shiners and other forage fishes

of Clear Lake, Iowa. Proceedings of the Iowa Academy of Science 70:215-223.

6. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

7. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

8. Peer, D.L. 1966. Relationship between size and maturity in the spottail shiner,

Notropis hudsonius. Journal of the Fisheries Research Board of Canada 23:455-

457.

9. Sewell, D.C. 1993. Food habits and distribution of the emerald shiner, Notropis

atherinoides Rafinesque, and the spottail shiner, Notropis hudsonius (Clinton), in

Lake Oahe, South Dakota. M.S. Thesis, South Dakota State University,

Brookings, South Dakota.

10. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press. 683 p. 300

11. Wells, L. 1980. Food of alewives, yellow perch, spottail shiners, trout-perch, and

slimy and fourhorn sculpins in southeastern Lake Michigan. Technical Papers of

the United States Department of the Interior Fish and Wildlife Service, 98.

12. Wells, L., and R. House. 1974. Life history of the spottail shiner (Notropis

hudsonius) in southeastern Lake Michigan, the Kalamazoo River, and western

Lake Erie. United States Department of the Interior Fish and Wildlife Service,

Research Report 78.

301

Carmine Shiner, Notropis percobromus (Cope, 1871)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; percobromus = unknown.

Description: Body fusiform, slender, elongate. Dorsally olive green with evenly distributed melanophores; dorsal scales outlined in slightly darker pigment creating a faint and less prominent “crosshatched” pattern; laterally silver to irredescent blue with a slightly faint rosy to pink lateral stripe extending onto opercle and cheek; ventrally silver to white; fins generally clear. Head moderate, slightly elongate. Snout blunt, conical, length slightly greater than diameter of eye. Eye large, placed laterally on head. Mouth large, terminal, slightly oblique with upper jaw extending to or slightly beyond anterior edge of eye. Barbels absent. Frenum absent. Lips pigmented, not extending onto chin.

Pharyngeal tooth pattern 2,4-4,2. Gill rakers short, 5-7. Dorsal fin with 8 rays and rounded tip; anterior fin ray does not extend past posterior fin ray when depressed.

Adipose fin absent. Caudal peduncle slender, slightly elongate. Caudal fin moderately forked. Anal fin with 9-10 rays and straight distal end. Pelvic fins abdominal with 8 rays; insertions distinctly anterior to insertion of dorsal fin. Pectoral fins with 12-13 rays.

Lateral line slightly decurved with 36-40 cycloid scales in series. Spawning males develop numerous minute tubercles on head and pectoral fin rays, as well as red, pink, or orange coloration on the head, cheeks, body, and base of the fins. Spawning females develop less intense coloration as well as minute and less numerous tubercles.

Similar Species: Closely resembles the Emerald Shiner. Emerald Shiner have a more bluntly rounded snout that is shorter than the diameter of the eye, 10-12 gill rakers and a dorsal fin with a pointed tip. Carmine Shiner are closely related to the Rosyface Shiner 302

(Notropis rubellus) and were previously referred to as Rosyface Shiner in South Dakota due to their morphological similarities.1,6 Recent studies have since determined the two species to be genetically distinct as they have geographically different distributions.6,12

Distribution and Habitat: Native to southern Manitoba, Canada, and the Midwestern

United States from eastern North and South Dakota, Minnesota and Wisconsin in the north, and south to Indiana, Arkansas, and Oklahoma.8 Native to the Sheyenne and Red

River drainages in North Dakota, as well as the Minnesota, James, and Big Sioux River drainages in eastern South Dakota.1,7 Have previously been reported from the Cannonball and Heart River watersheds of North Dakota. Information regarding required or preferred natural habitat of the species is limited or not well known as much of the literature pertains to studies conducted on Rosyface Shiners in areas outside of the Carmine Shiner distribution.3 The species currently occupies habitat spanning a wide range of climactic conditions.8 In the Minnesota drainage basin of South Dakota, Carmine Shiners are reported to be associated with the headwaters of creeks and small rivers with perennial flow, and no size discrepancy or confluence of a creek with a small river.6 In Manitoba,

Carmine Shiners are generally found in the mid-waters of fast-flowing creeks or small rivers with clear to semi-turbid waters and clean gravel or rubble substrate.3 They often occupy areas in or near riffles and pools, and may move into deeper areas such as pools during the winter.3,8 In aquaria with water temperatures ranging 15.5-31.4 °C (59.9-88.5

°F), mean minimum temperature was 17.8 °C (64.1 °F), mean maximum temperature was

28.8 °C (83.8 °F), and mean preferred temperature was 23.6 °C (74.5 °F).11

Reproduction: Little information is known about the species reproduction and spawning habits, although they are likely similar to the Rosyface Shiner.3 Minimum length of 303 sexually mature individuals from southern Manitoba reported as ≥46 mm (1.81 in) FL.4

Spawning individuals from Manitoba have been sampled in water temperatures 19.3-22.5

°C (66.7-72.5 °F) with water velocities up to 0.53 m/s, and at depths of 0.2-1.4 m (4.59 ft).3 In the more southern part of the species range, spawning occurs in water temperatures 20-28.9 °C (68-84.0 °F) with peak activity occurring from late April or early May through June.2,3,5,10 Spawning takes place in riffles over sand, cobble, boulder, or bedrock substrates.3 Fecundity of females from Manitoba ranged 694-2,806 eggs, with the number of eggs increasing with the size and age of the female.2,3 Eggs yellow in color, demersal, adhesive, and roughly 1.5 mm (0.06 in) in diameter.2,3,9 Hatching occurs within 57-59 hours at a water temperature of 21.2 °C (70.2 °F).2,3

Age and Growth: Length-at-age ranges from Wisconsin individuals are reported as: age-

1, 52-60 mm (2.05-2.36 in) TL; age-2, 58-75 mm (2.28-2.95 in) TL; age-3, 69-76 mm

(2.72-2.99 in) TL.2 Capable of reaching 88.9 mm (3.5 in).10 Longevity 3 years.2

Food and Feeding: Benthic, visual omnivore.3,9 Consume a variety of immature and mature aquatic and terrestrial invertebrates, especially Dipterans during the summer.3

Young-of-the-year primarily feed on diatoms and algae, consuming more insects as they grow older.10

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 304

3. Carmine Shiner Recovery Team. 2007. Recovery strategy for the carmine shiner

(Notropis percobromus) in Canada [Proposed]. Species at Risk Act Recovery

Strategy Series, Fisheries and Oceans Canada, Ottawa.

4. Carr, M., D.A. Watkinson, J.C. Svedsen, E.C. Enders, J.M. Long, and K.E.

Lindenschmidt. 2015. Geospatial modeling of the Birch River: distribution of

carmine shiner (Notropis percobromus) in geomorphic response units (GRU).

International Review of Hydrobiology 100:129-140.

5. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

6. Hayer, C.A., S.S. Wall, and C.R. Berry, Jr. 2008. Evaluation of predicted fish

distribution models for rare fish species in South Dakota. North American Journal

of Fisheries Management 28:1259-1269.

7. Hoagstrom, C.W, S.S. Wall, J.G Kral, B.G. Blackwell, and C.R. Berry, Jr. 2006.

Zoogeographic patterns and faunal change of South Dakota fishes. Western North

American Naturalist 67:161-184.

8. Pandit, S.N., B.M. Maitland, L.K. Pandit, M.S. Poesch, and E.C. Enders. 2017.

Climate change risks, extinction debt, and conservation implications for a

threatened freshwater fish: carmine shiner (Notropis percobromus). Science of the

Total Environment 598:1-11.

9. Pfeiffer, R.A. 1955. Studies on the life history of the rosyface shiner, Notropis

rubellus. Copeia 1955:95-104.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 305

11. Stol, J.A., J.C. Svendsen, and E.C. Enders. 2013. Determining the thermal

preferences of Carmine Shiner (Notropis percobromus) and Lake Sturgeon

(Acipenser fulvescens) using an automated shuttlebox. Canadian Technical Report

of Fisheries and Aquatic Sciences 3038.

12. Wood, R.M., R.L. Mayden, R.H. Matson, B.R. Kuhajda and S.R. Laymen. 2002.

Systematics and biogeography of the Notropis rubellus species group (Teleostei:

Cyprinidae). Bulletin of the Alabama Museum of Natural History 22:37-80.

306

Silverband Shiner, Notropis shumardi (Girard, 1856)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; shumardi = named after George C. Shumard, a naturalist on the Mexican Boundary Survey.

Description: Body fusiform, slender, elongate. Dorsally tan to silver with scales faintly outlined in darker pigment; laterally silver with no distinct markings; ventrally silver to white; fins generally clear. Head small. Snout bluntly rounded, not protruding beyond the mouth. Eye moderately large, placed laterally on head. Mouth large, terminal and oblique; upper jaw extends to anterior end of eye. Barbels absent. Frenum absent.

Pharyngeal tooth pattern variable between 1,4-4,1 and 2,4-4,2. Gill rakers 7-9. Dorsal fin with 8 rays and pointed tip; distal end straight to slightly falcate. Adipose fin absent.

Caudal peduncle moderately short, slender. Caudal fin forked. Anal fin with 8-9 rays and straight distal end. Pelvic fins abdominal with 9 rays; insertions slightly posterior to insertion of dorsal fin. Pectoral fins with 13-16 rays. Lateral line complete with 34-37 cycloid scales in series. Spawning males develop small tubercles on the dorsal and lateral sides of the head, as well as on the nape and breast regions of the body and pectoral fin rays. Juveniles similar to adults.

Similar Species: Closely resembles the Emerald Shiner, Sand Shiner, Channel Shiner, and River Shiner. Emerald Shiner have an anal fin with 10-13 rays, and a dorsal fin insertion that is distinctly posterior to the insertion of the pelvic fins. Sand Shiner display small, paired, dark specks or “mouse tracks” outlining the lateral line and a faint, dusky spot sometimes present at the base of the caudal fin. Channel Shiner have 8 pelvic fin rays and a lateral stripe expanding ventrally at the base of the caudal fin. River Shiner have 8 pelvic fin rays and 7 anal fin rays. 307

Distribution and Habitat: Native to the Missouri and Mississippi River basins from

South Dakota in the north, east to the Illinois and lower Ohio Rivers, south to the

Arkansas and Red Rivers to the Gulf of Mexico, with several native isolated populations near the Gulf slope in Texas.5 Occurs in the White, Missouri, Vermillion and James

Rivers in South Dakota and has also been recorded from Fort Randall Reservoir, Lake

Francis Case, Lake Oahe, and Lewis and Clark Lake.1,3,4,10 Considered rare within South

Dakota, as Nebraska and South Dakota represent the northernmost part of the species range.1,6,7 Inhabits the main channel, slackwater pools downstream of pointbars, and embayment’s in large turbid rivers with moderate to strong current over sand, gravel, silt, shale or mud substrate.5,8

Reproduction: Little information is known about the spawning and reproductive habits of the Silverband Shiner, especially in its northern range. Spawning season is prolonged and is said to take place from late May through mid-August in Tennessee, and from June to August in Lousiana.2,9 Sexual maturity is reached at age-1.11 Fractional, broadcast spawners, with multiple clutches of eggs being produced throughout the season.9,11 Clutch sizes from females ranging 47-76 mm (1.85-2.99 in) TL in the Brazos River, Texas averaged 387.3 and ranged from 144-750.11 Clutch size has shown to be positively correlated to total length of the female.11

Age and Growth: Age-0 individuals exhibit rapid growth.11 Mean lengths-at-age of individuals from Texas are reported as: age-0, 41 mm (1.61 in) TL; age-1, 66.5 mm (2.62 in) TL; age-2, 86 mm (3.39 in) TL.11 Capable of reaching 91.44 mm (3.6 in) TL.8

Longevity 2.5 years.11 308

Food and Feeding: Invertivores.11 Little details are known about the diet and feeding habits of the Silverband Shiner. Diets primarily consist of aquatic insects such as

Trichoptera, Ephemeroptera, and Diptera, but also include smaller portions of detritus terrestrial insects, crustaceans, and algae.11 Aquatic insects become more important in the diet of adults.11

Literature Cited:

1. Berry, C.R. Jr., and B. Young. 2004. Fishes of the Missouri national recreational

river, South Dakota and Nebraska. Great Plains Research 14:89-114.

2. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

3. Harlan, J.R., and E.B. Speaker. 1987. Iowa fish and fishing. Des Moines: Iowa

Department of Natural Resources.

4. Jones, S.J. 2018. Western prairie stream fisheries: an assessment of past and

present fish assemblage structure, biotic homogenization, and population

dynamics in western South Dakota streams. M.S. Thesis, South Dakota State

University, Brookings, South Dakota.

5. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

6. Madsen, T.I. 1985. The status and distribution of the uncommon fishes in

Nebraska. M.S. Thesis, University of Nebraska-Omaha.

7. Miller, R.R. 1972. Threatened freshwater fishes of the United States. Transactions

of the American Fisheries Society 101:239-252. 309

8. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of

Conservation, Jefferson City, Missouri.

9. Suttkus, R.D. 1980. Notropis candidus, a new cyprinid fish from the Mobile Bay

basin, and a review of the nomenclatural history of Notropis shumardi (Girard).

Bulletin of the Alabama Museum of Natural History 5:1-15.

10. Underhill, J.C. 1959. Fishes of the Vermillion River, South Dakota. Proceedings

of the South Dakota Academy of Science 38:96-102.

11. Williams, C.S. 2011. Life history characteristics of three obligate riverine species

and drift patterns of lower Brazos River fishes. Pd.D. Dissertation, Texas State

University, San Marcos, Texas.

310

Sand Shiner, Notropis stramineus (Cope, 1865)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; stramineus = Latin for “made of straw”, likely referring to the species coloration.

Description: Body fusiform, slender, laterally compressed. Dorsally light olive to tan with dorsal scales outlined in darker pigment creating a “crosshatched” pattern; prominent, narrow mid-dorsal stripe present that widens into a wedge-shaped spot at the anterior base of the dorsal fin and does not surround the fin base; laterally silver with small, paired, dark specks or “mouse tracks” outlining the lateral line; lateral stripe faint, continuous from caudal peduncle to eye, but fading anterior to dorsal fin; ventrally silver to white; fins with little to no pigment; faint, dusky spot sometimes present at base of caudal fin. Head short. Snout blunt, rounded; does not protrude past the mouth. Eye large, placed laterally on head. Mouth small, subterminal and slightly oblique with lower jaw extending to anterior edge of the eye; length of upper jaw less than the diameter of the eye. Barbels absent. Frenum absent. Pharyngeal teeth 0,4-4,0. Gill rakers short, roughly

8. Dorsal fin with 8 rays. Adipose fin absent. Caudal peduncle slender, slightly elongate.

Caudal fin moderately forked. Anal fin with 7 rays and straight distal end. Pelvic fins abdominal with 8 rays, insertions directly under or slightly anterior to insertion of dorsal fin. Pectoral fins with 13-15 rays. Lateral line complete, slightly decurved with 31-35 cycloid scales in series. Spawning males develop small tubercles on the dorsal and lateral sides of the head, as well as on the pectoral fin rays. Juveniles similar to adults.

Similar Species: Closely resembles the Topeka Shiner, Bigmouth Shiner, and River

Shiner. Topeka Shiner tend to have a deeper body, a nearly straight lateral line, a triangular or chevron shaped black mark at the base of the caudal fin, and spawning 311 males with red-orange fins and sides of head. Bigmouth Shiner with a mid-dorsal stripe that does not widen at the base of the dorsal fin but instead surrounds it, 8 anal fin rays, a snout that slightly overhangs the mouth, and eyes directed upwards with lower margins of pupils visible when fish is viewed dorsally. River Shiner lack any small, paired, dark specks or “mouse tracks” outlining the lateral line, have a larger mouth, and a mid-dorsal stripe that does not widen at the base of the dorsal fin but instead surrounds it.

Distribution and Habitat: Native throughout much of central North America from the headwaters of the Platte River in eastern Wyoming, north throughout the Missouri, lower

Red River of the North drainage in Canada, Mississippi and St. Lawrence-Great Lakes drainages, east to the Tennessee River drainage, and south to the Rio Grande River drainage in New Mexico and Texas.3 Vastly distributed across North and South Dakota.

Inhabits a wide variety of habitats including lakes, ponds, rivers, streams and creeks.1,6

Prefers to school in the benthic or near benthic environments of pools with considerable to moderate velocities as well as sand or gravel riffles in moderate to large sized streams of moderate to high gradient with clear to slightly turbid water.9,13 Abundance increases in areas with higher amounts of sand substrate.5 Laboratory experiments have shown that the Sand Shiners habitat preferences are known to be influenced by interactions within a fish assemblage, choosing no flow areas when only Sand Shiner are present, and high flow areas in mixed species groups.5 Absent or rare in low gradient or intermittent waters, or in areas with increased levels of aquatic vegetation, erosion and siltation.5,13

Reproduction: Little information is known about the spawning and reproductive habits of Sand Shiner. Spawning takes place May through August, and peaks when water temperatures reach 27-37 °C (80.6-98.6 °F).11,12 Sexual maturity reached at age-1, or 312 when fish average 49.1 mm (1.93 in) TL, with mortality increasing in age-2 fish following their second year of spawning.12 In aquaria, a single male has been observed chasing a single female prior to spawning, but little is known about the actual spawning event or spawning behavior.7 Spawning known to take place over clean gravel and sand substrate.4 Broadcast spawner.7 Fecundity tends to increase with the size of the female, as females from Iowa contained roughly 250 eggs age age-1, 1,100 eggs at age-2, and 1,800 eggs at age-3.11 Eggs roughly 0.65-0.75 mm (0.02-0.03 in) in diameter, demersal, adhesive, and quickly settle into the substrate.7,12

Age and Growth: Mean lengths-at-age of individuals from Iowa are reported as: age-1,

38.5 mm (1.52 in) TL; age-2, 53.0 mm (2.09 in) TL; age-3, 63.9 mm (2.52 in) TL.8

Capable of reaching 87 mm (3.43 in) TL.8 Longevity 3 years.8

Food and Feeding: Generalized, opportunistic, benthic omnivores.2,10 Known to actively feed throughout the day and are relatively inactive at night.10 Primarily consumes a wide variety of immature and adult aquatic and terrestrial insects such as small mayfly nymphs, stoneflies, and midge larvae.2 Also known to consume detritus, and plant material.10

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Gillen, A.L., and T. Hart. 1980. Feeding interrelationships between the sand

shiner and the striped shiner. Ohio Journal of Science 80:71-76.

3. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp. 313

4. Miller, R.J., and H.W. Robinson. 2004. Fishes of Oklahoma, revised edition.

University of Oklahoma Press, Norman, Oklahoma.

5. Mueller, R. Jr., and M. Pyron. 2009. Substrate and current velocity preferences of

spotfin shiner (Cyprinella spiloptera) and sand shiner (Notropis stramineus) in

artificial streams. Journal of Freshwater Ecology 24:239-245.

6. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

7. Platania, S.P., and C.S. Altenbach. 1998. Reproductive strategies and egg types of

seven Rio Grande basin cyprinids. Copeia 1998: 559-569.

8. Smith, C.D., T.E. Neebling, and M.C. Quist. 2010. Population dynamics of the

sand shiner (Notropis stramineus) in non-wadeable rivers of Iowa. Journal of

Freshwater Ecology 25:617-626.

9. Smith, P.W. 1979. The fishes of Illinois. University of Illinois Press, Urbana,

Illinois.

10. Starrett, W.C. 1950. Food relationships of the minnows of the Des Moines River,

Iowa. Ecology 31:216-233.

11. Starrett, W.C. 1951. Some factors affecting the abundance of minnows in the Des

Moines River, Iowa. Ecology 32:13-27.

12. Summerfelt, R.C., and C.O. Minckley. 1969. Aspects of the life history of the

sand shiner, Notropis stramineus (Cope), in the Smoky Hill River, Kansas.

Transactions of the American Fisheries Society 98:444-453.

13. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press. 683 p.

314

Topeka Shiner, Notropis topeka (Gilbert, 1884)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; topeka = referring to

Topeka, Kansas, the type locality.

Description: Body fusiform, moderately deep, laterally compressed. Dorsally light olive to tan with dorsal scales outlined in darker pigment creating a “crosshatched” pattern; prominent, narrow mid-dorsal stripe present that slightly widens at the base of the dorsal fin and does not surround the fin base; laterally silver with small, paired, dark specks or

“mouse tracks” outlining the lateral line; lateral stripe dusky, extending from caudal peduncle to snout; ventrally silver to white; fins with little to no pigment; triangular/ chevron shaped black spot present at the base of the caudal fin. Head short. Snout blunt, rounded. Eye large, placed laterally on head. Mouth small, terminal to slightly subterminal and oblique with lower jaw slightly extending beyond anterior edge of eye; length of upper jaw less than the diameter of the eye. Barbels absent. Frenum absent.

Pharyngeal tooth pattern 0,4-4,0. Dorsal fin with 8 rays. Adipose fin absent. Caudal peduncle slender, slightly elongate. Caudal fin moderately forked. Anal fin with 7 rays and straight distal end. Pelvic fins abdominal with 8 rays, insertions directly under or slightly anterior to insertion of dorsal fin. Lateral line complete, nearly straight with 32-

37 cycloid scales in series. Spawning males with bright red-orange coloration on fins and side of head, a faint blue hue or luminance on lateral sides, and small tubercles on the snout, head and body

Similar Species: Closely resembles the Sand Shiner, Bigmouth Shiner, and River Shiner.

Sand Shiner tend to have less deep of a body, a slightly decurved lateral line, and a faint, dusky spot sometimes present at the base of the caudal fin. Spawning male Sand Shiners 315 do not develop any red-orange coloration on fins or side of head. Bigmouth Shiner with a mid-dorsal stripe that does not widen at the base of the dorsal fin but instead surrounds it,

8 anal fin rays, a snout that slightly overhangs the mouth, and eyes directed upwards with lower margins of pupils visible when fish is viewed from above. River Shiner lack any small, paired, dark specks or “mouse tracks” outlining the lateral line, have a larger mouth, and a mid-dorsal stripe that does not widen at the base of the dorsal fin but instead surrounds it.

Distribution and Habitat: Native range extends throughout parts of the Mississippi,

Missouri, and Arkansas river drainages from central Missouri and Kansas to southern

Minnesota, eastern South Dakota, Iowa and Nebraska.2 Once widespread and abundant throughout the central prairie region of the United States, the species is now found only in highly fragmented populations which make up less than 10% of their historical range.4,15 While they are not found in abundant numbers, Topeka Shiner are widespread throughout the Big Sioux, Vermillion, and James river basins in eastern South Dakota.4,13

Recent studies have suggested that Topeka Shiners are more abundant in South Dakota than any other state within the species range.13 The decline of the species historical distribution and abundance has been attributed to extensive changes in stream hydrology caused by increased sedimentation and reduced water quality from agricultural practices, non-point source pollution, hydrologic changes from the construction of impoundments, and increased predation by non-native stocked species.1,4,11,16 Inhabit small, quiet, clear, and moderately cool low-order prairie and upland creeks, as well as intermittent streams, tributaries and headwater reaches of larger streams in off-channel habitats with sand, gravel or cobble substrate and submerged vegetation.3,4,5,12 Often found within the 316 limnetic zone of large pool and slackwater habitats avoiding water velocities >10 cm/s, however they are capable of swimming faster speeds than water velocities in habitats they normally utilize.1,4,12 The majority of sites where Topeka Shiner were found in eastern South Dakota had water velocities <0.20 m/s.4 Paired with the species swimming ability, fish ways and culverts may help facilitate dispersal and recolonization of the species.1 Avoids high water velocities as they can reduce population size and negatively impact reproduction.1,11 Tolerant of high water temperatures with a critical thermal maximum reported as 39 °C (102.2 °F) when acclimated to water 31 °C (87.8 °F).10

Capable of tolerating oxygen concentrations as low as 1.2 mg/L.10 Juveniles occupy shallow off-channel habitat.3

Reproduction: Little information is known about the Topeka Shiners reproductive habits. Spawning takes place from mid-May to early August with peak spawning occurring in June.5 In aquaria, spawning has been observed to begin when water temperature reaches 24.4 °C (75.92 °F).8 Sexual maturity begins at age-1, with most individuals maturing at age-2.5,9 Mature males are territorial and defend small areas <0.5 m (1.64 ft) in diameter on the edge of Green Sunfish nests.5,8,9,12,14 In Missouri, Topeka

Shiner have been known to spawn over Green Sunfish and Orangespotted Sunfish nests.9,12 Spawning behavior consists of males swimming parallel to females, and vibrating their bodies together to initiate the release of eggs and milt which are broascast.8,14 Females repeat the spawning process two to four times before they resume normal swimming activity.8 Mean clutch size of eggs-at-age of 66 mature females from

Minnesota are reported as: age-1, 351 mature ova; age-2, 559 mature ova; age-3, 478 mature ova.5 Eggs yellow to yellow-orange in color, adhesive, and roughly 0.73-0.97 mm 317

(0.03-0.04 in) in diameter.5,8,9 Clutch size is more correlated with size of the female than age.5 Hatching occurs in 5 days at 22.2 °C (72 °F).8

Age and Growth: Majority of age-1 and older individuals achieved most of their annual growth from May to July.5 Optimal growth temperature 27 °C (80.6 °F).10 Males generally achieve longer lengths-at-age than females.5 Mean lengths-at-age of males from southwestern Minnesota are reported as: age-1, 29.6 mm (1.17 in) SL; age-2, 46.3 mm

(1.82 in) SL; age-3, 55.6 mm (2.12 in) SL.5 Mean lengths-at-age of females reported as: age-1, 29.3 mm (1.15 in) SL; age-2, 41.1 mm (1.62 in) SL; age-3, 46.5 mm (1.83 in) SL.5

Ages 1-3 are known to experience high mortality rates towards the end of the summer, prior to the recruitment of young-of-the-years.5 Capable of reaching 75 mm (2.95 in) TL.4

Longevity 3 years.5,12

Food and Feeding: Generalized, opportunistic, benthic and nektonic omnivores.5,6,7

Forage diurnally with peak feeding activity occurring from the early morning to early afternoon.9 Adult diets mainly consist of microcrustaceans and adult and larval stages of aquatic and terrestrial insects such as Diptera, Ephemeroptera, and Coleoptera.5,9 Larval fish, algae, snails, worms, vascular plants, and detritus have also been found in diets.5,7

Literature Cited:

1. Adams, S.R., J.J. Hover, and K.J. Killgore. 2000. Swimming performance of the

Topeka shiner (Notropis topeka) an endangered Midwestern minnow. The

American Midland Naturalist 144:178-186.

2. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor. 318

3. Bakevich, B.D., C.L. Pierce, and M.C. Quist. 2013. Habitat, fish species, and fish

assemblage associations of the Topeka shiner in west-central Iowa. North

American Journal of Fisheries Management 33:1258-1268.

4. Blausey, C.M. 2001. The status and distribution of the Topeka shiner Notropis

topeka in eastern South Dakota. M.S. Thesis, South Dakota State University,

Brookings, South Dakota.

5. Dahle, S.P. 2001. Studies of Topeka shiner (Notropis topeka) life history and

distribution in Minnesota. M.S. Thesis, University of Minnesota, Minneapolis,

Minnesota.

6. Hatch, J.T., and S. Besaw 1998. Diverse food use in Minnesota populations of the

Topeka shiner (Notropis topeka). General College and James Ford Bell Museum

of Natural History. University of Minnesota, Minneapolis.

7. Hatch, J.T., and S. Besaw. 2001. Food use in Minnesota populations of the

Topeka shiner (Notropis topeka). Journal of Freshwater Ecology 16:229-233.

8. Katula, R. 1998. Eureka Topeka! Tropical Fish Hobbyist 47:54-60.

9. Kerns, H.A., and J.L. Bonneau. 2002. Aspects of the life history and feeding

habits of the Topeka shiner (Notropis topeka) in Kansas. Transactions of the

Kansas Academy of Science 105:125-142.

10. Koehle, J.J., and I.R. Adelman, 2007. The effects of temperature, dissolved

oxygen, and Asian tapeworm infection on growth and survival of the Topeka

Shiner. Transactions of the American Fisheries Society 136:1607-1613. 319

11. Minckley, W.L., and F.B. Cross. 1959. Distribution, habitat, and abundance of the

Topeka shiner Notropis topeka (Gilbert) in Kansas. The American Midland

Naturalist 61:210-217.

12. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

13. Shearer, J.S. 2003. Topeka shiner (Notropis topeka) management plan for the

state of South Dakota. South Dakota Department of Game, Fish and Parks, Pierre,

Wildlife Division Report No. 2003-10, 82pp.

14. Stark, W. J., J. S. Luginbill, and M. E. Erbele. 2002. Natural history of a relict

population of Topeka shiner (Notropis topeka) in northwestern Kansas.

Transactions of the Kansas Academy of Science 105:143-152.

15. Tabor, V.M. 1998. Final rule to list the Topeka shiner as endangered. Federal

Register. 63:69008-69021.

16. USFWS. 1998. Endangered and threatened wildlife and plants; final rule to list

the Topeka Shiner as endangered. Federal Register 63:204(15 December

1998):69008-69021.

320

Channel Shiner, Notropis wickliffi (Trautman, 1931)

Etymology and Synonyms: Notropis = Greek for “back” or “keel”; wickliffi = named after Ohio fishes student Edward L. Wickliff, also a friend of Dr. Milton B. Trautman.

Description: Body fusiform, slightly deep, laterally compressed. Dorsally dusky gray to olive with scales outlined in darker pigment creating a prominent “crosshatched” pattern; mid-dorsal stripe faint to absent; laterally silver with small, paired, faint, darker specks or

“mouse tracks” outlining the lateral line; lateral stripe absent to faint, expanding ventrally at the base of the caudal fin to form a wedge; ventrally silver to white; fins clear to lightly pigmented. Head elongate. Snout bluntly rounded. Eye large, placed laterally on head; diameter greater than snout length. Mouth small, terminal to slightly subterminal and oblique; upper jaw extending to anterior edge of eye. Frenum absent. Barbels absent.

Pharyngeal tooth pattern 0,4-4,0. Gill rakers short, roughly 5. Dorsal fin with 8 rays, straight distal end and pointed tip. Adipose fin absent. Caudal peduncle slender, slightly elongate. Caudal fin moderately forked. Anal fin with 8 rays and straight distal end.

Pelvic fins abdominal with 8 rays; insertions inferior or slightly anterior to insertion of dorsal fin. Pectoral fins with 14-16 rays. Lateral line complete with 32-39 cycloid scales in series. Spawning males develop small tubercles on the head and pectoral fins.

Juveniles similar to adults.

Similar Species: Recent taxonomic confusion with the Mimic Shiner, Notropis volucellus, is the main reason for scarce knowledge of the Channel Shiners life history traits and questionable geographic distribution.1 In the Dakotas, the species may be confused with the Bigmouth Shiner, Emerald Shiner, River Shiner and Sand Shiner.

Bigmouth Shiner have a large, slightly subterminal mouth with the lower jaw extending 321 to the middle of the eye, and the length of the upper jaw greater than diameter of the eye.

Emerald Shiner have an anal fin with 10-13 rays, and a dorsal fin insertion that is distinctly posterior to the insertion of the pelvic fins. River Shiner have a dark, well- defined mid-dorsal stripe present that surrounds the dorsal fin base, and an upper jaw that extends to or slightly beyond anterior end of eye and is greater in length than the diameter of the eye. Sand Shiner have an anal fin with 7 rays, and an upper jaw that is shorter than the diameter of the eye.

Distribution and Habitat: Due to the confusion with the Mimic Shiner, the native distribution of Channel Shiner is rather limited and poorly understood, but is known to include the Mississippi River and its larger tributaries.2,4 Documented in the Wild Rice

River within the Red River of the North watershed in North Dakota. Recorded only from the Missouri River near Yankton in southeastern South Dakota. Inhabits deep, midwaters or pools in areas with little to no current with silt to gravel substrates in large rivers and the lower portions of their larger tributaries.2,4 The species is thought to be fairly tolerant of turbidity due to it being collected from an array of substrate types.4

Reproduction: Information on spawning and reproductive habits of the Channel Shiner is extremely limited, especially within the upper Mississippi River basin near the

Dakotas. In Ohio, spawning is known to take place from June to August.4 In Tennessee, breeding tubercles have not been observed on individuals <50 mm (1.97 in) TL.2 Eggs are broadcast over sand or gravel substrate.3 No nest is constructed and no parental care is given.3

Age and Growth: Little information is available on the growth of Channel Shiner, especially around the Dakotas region. Young-of-year from Ohio sampled in October 322 measured 20-41 mm (0.79-1.61 in) in length, and 30-64 mm (1.18-2.52 in) in length around age-1.4 Capable of reaching 78 mm (3.07 in) TL.2

Food and Feeding: The diet and feeding habits of the Channel Shiner remain unstudied, but they are likely to consume various aquatic invertebrates.3

Literature Cited:

1. Dowell, S.A. 2010. Morphological and genetic investigations of Pennsylvania

populations of the channel shiner, Notropis wickliffi. M.S. Thesis, Duquesne

University, Pittsburgh, Pennsylvania.

2. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

3. Ohio Department of Natural Resources. 2012. Ohio Department of Natural

Resources Division of Wildlife, Channel Shiner – Notropis wickliffi,

http://wildlife.ohiodnr.gov/species-and-habitats/species-guide-index/fish/channel-

shiner.

4. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

323

Suckermouth Minnow, Phenacobius mirabilis (Girard, 1856)

Etymology and Synonyms: Phenacobius = Greek for “deceptive life”, likely referring to the suggested herbivorous lifestyle due to a long intestine, but it is not a herbivorous species; mirabilis = Latin for “strange” or “wonderful”.

Description: Body fusiform, slender, elongate, cylindrical anteriorly, slightly laterally compressed posteriorly. Dorsally dark gray to olive; laterally silver to tan or light olive with dark, dusky lateral stripe ending in a prominent, dark spot at the base of the caudal fin; scales above lateral stripe outlined in darker pigment creating a herringbone or crosshatched pattern; ventrally silver to white; pectoral, dorsal, and caudal fins lightly pigmented with olive to tan coloration; anal and pelvic fins typically white to clear. Head short, slightly dorsoventrally compressed. Snout elongate, bluntly rounded, protruding beyond mouth. Eye moderately large, placed laterally on upper portion of head. Mouth small, subterminal, sucker-like; upper jaw does not extend to anterior edge of eye.

Frenum absent. Barbels absent. Lips fleshy; lower lip with a fleshy posterior lobe on each side. Pharyngeal tooth pattern 0,4-4,0. Gill rakers short, 9. Dorsal fin with 8 rays and straight to slightly concave distal end. Adipose fin absent. Caudal peduncle moderately short, uniform in thickness. Caudal fin moderately forked. Anal fin with 6-7 rays and straight distal end. Pelvic fins abdominal with 8 rays; insertions slightly posterior to insertion of dorsal fin. Pectoral fins with 14-16 rays. Lateral line complete, slightly decurved with 40-51 cycloid scales in series. Predorsal scale rows typically 18-21.

Spawning males develop small tubercles on dorsal side of head, anterior end of body, and pectoral fins. Females may develop small, less numerous tubercles on the dorsal side of head. 324

Similar Species: Closely resembles the Longnose Dace, Western Blacknose Dace,

Central Stoneroller and Shoal Chub. Both Longnose Dace and Western Blacknose Dace have a frenum and barbels present, pelvic fin insertions slightly anterior to the insertion of the dorsal fin, and >60 small cycloid scales in lateral line series. Central Stoneroller have a more robust body with mottling on the lateral sides, and a cartilaginous ridge on the inner lower lip. Shoal Chub have small, prominent, black, scattered specks on the dorsal and lateral sides with barbels present.

Distribution and Habitat: Native to the plains ecoregions of the United States ranging from South Dakota, eastern Wyoming, and Colorado in the west, east throughout the

Mississippi River basin to Ohio, and south throughout the Tennessee river valleys to

Texas and northeastern New Mexico.6 The species limited distribution and abundance is likely due to habitat alteration, fragmentation, groundwater withdrawal and impoundments.3 Frequent in small to large streams of low to high gradient with permanent flow, deep pools, riffles and runs over gravel substrate.3,7 The species appears to be tolerant of fluctuating water and higher turbidity levels as long as the gradient is sufficient and there is continuous flow to keep the gravel substrate free of silt.7,8 Young- of-the-year inhabit riffles over gravel substrate like adults, but are also frequent in deep, low velocity pools or backwaters over sand and silt substrate.3,5

Reproduction: Spawning season is prolonged and takes place March through August.3,4

Sexual maturity occurs at age-2.1 Fractional spawners, with females likely spawning two or more times a season and releasing only a small batch of ripe eggs at a time.1,2 Smaller females spawn earlier than larger females.5 It is suggested that spawning takes place within the species year-round habitat consisting of riffles with gravel or cobble substrate, 325 however they have also been observed spawning in shallow microhabitats and migrate to deeper depths postspawning.2,4,5,7 In a laboratory setting, spawning was successful in water temperatures 17-23 °C (62.6-73.4 °F).3 In Missouri, some individuals began spawning when the water temperature reached 9 °C (48.2 °F), with peak spawning occurring at 12-20 °C (53.6-68 °F).4 Spawning behavior consists of individuals rapidly chasing each other in circles near the substrate, and males positioning themselves perpendicular to the female, nudging their vent areas, and rapidly shuddering to likely initiate the release of eggs and milt.2,3 Fecundity roughly 200-500 eggs per female, with larger females generally producing a greater number of eggs than smaller females.2,5 A 90 mm (3.54 in) TL female from Wisconsin produced approximately 1,640 eggs.1 Eggs adhesive, spherical, dark yellow to gold in color, and range 1.55-1.70 mm (0.06-0.07 in) in diameter one hour post fertilization.2,3 In aquaria, hatching begins at approximately 3 days at 23 °C (73.4 °F), and at 4 days in water temperatures 17-19 °C (62.6-66.2 °F).3

Age and Growth: Embryo development and growth of larvae are much faster at warmer water temperatures.3 Larvae average 4.6 mm (0.18 in) TL at hatching.2 In aquaria, larvae

10 days post-hatch held at 23 °C (73.4 °F) grew 0.54 mm/d (0.02 in/d), and 0.39 mm/d

(0.01 in/d) at 17 °C (62.6 °F).2,3 Up to 72% of their maximum length is achieved during the first growing season.5 Capable of reaching 122 mm (4.80 in).8 Longevity 3 years.5

Food and Feeding: Benthic insectivore. Primarily feeds on aquatic insects such as

Diptera, Trichoptera, and Ephemeroptera.5 It has been suggested that the species is a selective and sensitive feeder, using their numerous sensory organs and fleshy lips to feed by taste and touch rather than by sight.3,5

Literature Cited: 326

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Bestgen, K.R., and R.I. Compton. 2007. Reproduction and culture of suckermouth

minnow. North American Journal of Aquaculture 69:345-350.

3. Bestgen, K.R., K. Zelasko, and R. Compton. 2003. Environmental factors limiting

suckermouth minnow Phenacobius mirabilis populations in Colorado. Final report

submitted to the Colorado Division of Wildlife, Fort Collins. Larval Fish

Laboratory Contribution 136.

4. Brewer, S.K., D.M. Papoulias, and C.F. Rabeni. 2006. Spawning habitat

associations and selection by fishes in a flow-regulated prairie river. Transactions

of the American Fisheries Society 135:763-778.

5. Haas, M.A. 1977. Some aspects of the life history of the suckermouth minnow,

Phenacobius mirabilis (Girard). M.S. Thesis, University of Missouri, Columbia,

Missouri.

6. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication. 1980-12. 854pp.

7. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of

Conservation, Jefferson City.

8. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

327

Bluntnose Minnow, Pimephales notatus (Rafinesque, 1820)

Etymology and Synonyms: Pimephales = Greek for “fat head”; notatus = “marked”, or

“spotted”.

Description: Body fusiform, robust, rather cylindrical anteriorly and laterally compressed posteriorly. Dorsally dark olive to brown; laterally brown to tan with a dark lateral stripe extending onto the opercle and snout, and leading into a caudal spot that tends to fade with age; ventrally cream to white; fins lightly pigmented with small, dusky, dark spot sometimes present on larger individuals at the anterior base of the dorsal fin.

Head broad. Snout bluntly rounded, slightly protruding beyond the mouth. Eye moderately large, placed laterally on head. Mouth subterminal, moderately large; upper jaw extends to anterior edge of eye. Barbels absent. Frenum absent. Lips thin, uniform in thickness. Pharyngeal tooth pattern 0,4-4,0. Gill rakers short, 7-10. Dorsal fin with 8 rays and rounded distal end. Adipose fin absent. Caudal peduncle slightly elongate, thick.

Caudal fin moderately forked. Anal fin with 7-8 rays. Pelvic fins abdominal with 8 rays; insertions slightly anterior to insertion of dorsal fin. Pectoral fins with 15-16 rays. Lateral line complete with 37-44 cycloid scales in series. Predorsal scale rows generally 22 or fewer. Spawning males turn dark gray to black in color, and develop three rows of white, sharp tubercles on the snout and a single pair of small, flap-like barbels, one on each corner of the mouth. A patch of swollen, spongy skin called a “dorsal pad” is developed anterior to the dorsal fin, and the first ray of the dorsal fin is short and thickened.

Similar Species: Closely resembles the Fathead Minnow. Fathead Minnows generally have an incomplete lateral line, scales above the lateral line outlined in darker pigment creating a crosshatched or herringbone pattern, a smaller mouth with an upper jaw that 328 does not extend to the anterior edge of the eye, and no spot present at the base of the caudal fin. Brassy Minnow have a small mouth with an upper jaw that does not extend to the anterior edge of the eye.

Distribution and Habitat: Widespread native distribution throughout southern Canada from Manitoba to Quebec, and the northeast and central United States from the Dakotas south to Oklahoma in the west, east throughout the Mississippi, St. Lawrence-Great

Lakes, and Ohio River drainages to New York, south to Tennessee and the Gulf of

Mexico, but not occupying the Atlantic Slope drainages. Recorded from the James,

Sheyenne, and Red River basins of eastern North Dakota, but disjunct populations are also present in the Little Missouri, and Missouri River basins, north of Lake Sakakawea.

Primarily occurs throughout the Missouri, James, Vermillion, Big Sioux and Minnesota

River drainages in South Dakota, but have also been recorded west of the Missouri River from the Cheyenne and Belle Fourche drainages. Inhabits a wide variety of habitats including glacial lakes and reservoirs, but prefers quiet backwaters and pools of low gradient, medium to moderately large streams with sand or gravel substrate, clear to slightly turbid water, persistent flow and moderate amounts of submerged aquatic vegetation.1,2,8 Presence of populations has been shown to increase with an increase in floating overhead cover.5 Mean critical water temperature 36.6 °C (97.8 °F).12 Mean critical dissolved oxygen concentration reported as 1.04 mg/L.12 Prefers waters with a pH

≥ 7.8, but have been present in waters with pH values ≥ 5.6.6,10

Reproduction: Spawning occurs May through August, and begins when water temperatures reach roughly 21 °C (69.8 °F).3,7,9 Age at sexual maturity may vary between populations, but most often occurs at age 1-2.14 Nests are cavity-like and are constructed 329 under rocks, logs, or other structures along the bottom substrate, with eggs deposited in a single layer and adhering to the ceiling of the structure.9,11,14 Nests are generally found at depths up to 1 m (3.28 ft), but they may be as deep as 2.4 m (7.87 ft).9,14 If necessary, the male may dig out sediments himself.9 Females are known to prefer spawning with larger males, likely increasing the safety of their young.9 Spawning generally takes place at night.4,14 Fractional spawners, with time between spawning events ranging 2-14 days.4

Females may spawn >10 times during a single season.9,4 Eleven pairs of Bluntnose

Minnows from Pennsylvania produced 7-19 clutches of eggs during the spawning season, with the average clutch size ranging 93-239 eggs, to produce a total of 1,112-4,195 eggs per spawning season.4 The first clutch of eggs generally contains fewer eggs than later clutches.4 Fecundity is known to be unrelated to the length and weight of the female.4

Eggs roughly 1.45 mm (0.06 in) in diameter.14 Males provide parental care by guarding and cleaning the eggs until hatching.9 To help keep the eggs free of silt, snails or parasites, males clean the eggs by rubbing them with their dorsal pad.9 Hatching occurs within 6-7 days.9

Age and Growth: Larvae roughly 5 mm (0.20 in) TL one day post-hatching.14 Sexually dimorphic species, with males growing faster and reaching considerably larger lengths than females.7,8 Capable of reaching 0.17 mm (4.3 in) TL.13 Longevity 3 years.7

Food and Feeding: Benthic opportunistic omnivore. Often forage throughout the day along the bottom in large schools and consume a mixture of algae, large diatoms, aquatic insect larvae and microcrustacea.2,7 As individuals increase in size, the tendency for them to feed among aquatic vegetation and along the bottom substrate likely increases.7 330

Females are known to consume embryos in the nests when males are defending the nest from other males and small fish, or are out chasing potential females.9

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Carlander, K. 1969. Handbook of freshwater fishery biology, volume 1. Life

history data on freshwater fishes of the United States and Canada exclusive of the

Perciformes. Iowa State University Press, Ames, Iowa, USA.

4. Gale, W.F. 1983. Fecundity and spawning frequency of caged bluntnose

minnows-fractional spawners. Transactions of the American Fisheries Society

112:398-402.

5. Gatz, A.J. Jr. 2008. The use of floating overhead cover by warmwater stream

fishes. Hydrobiologia 600:307-310.

6. Matuszek, J.E., J. Goodier, and D.L. Wales. 1990. The occurrence of cyprinidae

and other small fish species in relation to pH in Ontario lakes. Transactions of the

American Fisheries Society 119:850-861.

7. Moyle, P.B. 1973. Ecological segregation among three species of minnows

(Cyprinidae) in a Minnesota lake. Transactions of the American Fisheries Society

102:794-805.

8. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 331

9. Pot, W. 1985. Competition for nests, parental care, and female choice in the

bluntnose minnow, Pimephales notatus. Ph.D. Dissertation, University of Guelph,

Guelph, Ontario, Canada.

10. Rahel, F.J., and J.J. Magnuson. 1983. Low pH and the absence of fish species in

naturally acidic Wisconsin lakes: inferences for cultural acidification. Canadian

Journal of Fisheries and Aquatic Sciences 40:3-9.

11. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

12. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

13. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press. 683 p.

14. Westman, J. 1938. Studies on the reproduction and growth of the bluntnose

minnow, Hyborhynchus notatus (Rafinesque). Copeia 1938:57-61.

332

Fathead Minnow, Pimephales promelas (Rafinesque, 1820)

Etymology and Synonyms: Pimephales = Greek for “fat head”; promelas = Greek for

“before black”, likely referring to the dark coloration on the head of spawning males.

Description: Body fusiform, robust, moderately deep. Dorsally dark olive to brown; laterally tan, olive to silver with a dark lateral stripe that tends to fade with age and absent caudal spot; scales above lateral line outlined in darker pigment creating a crosshatched or herringbone pattern; ventrally cream to white; fins lightly pigmented with small, dusky, dark spot sometimes present on larger individuals at the anterior base of the dorsal fin. Head broad. Snout bluntly rounded, not extending beyond the upper lip. Eye moderate, placed laterally on head. Mouth small, terminal and oblique; upper jaw does not extend to anterior edge of the eye. Barbels absent. Frenum absent. Lips thin.

Pharyngeal tooth pattern 0,4-4,0. Gill rakers, 14-16. Dorsal fin with 8 rays and rounded distal end. Adipose fin absent. Caudal peduncle slightly elongate, thick. Caudal fin moderately forked. Anal fin with 7 rays. Pelvic fins abdominal with 8 rays; insertions slightly anterior to insertion of dorsal fin. Pectoral fins with 15-16 rays. Lateral line incomplete with 40-54 cycloid scales in series. Predorsal scale rows generally 23 or more. Spawning males are often darker in color especially on the head. Males also develop large breeding tubercles on the head, snout and pectoral fins rays as well as a fleshy dorsal pad, and thickened, shorter anterior dorsal fin ray.

Similar Species: Closely resembles the Bluntnose Minnow. Bluntnose Minnow have a complete lateral line with scales above the lateral line not outlined in darker pigment.

Bluntnose Minnow also have a small subterminal mouth with an upper jaw that extends to the anterior edge of the eye, and a caudal spot typically present. Brassy Minnow have a 333 complete lateral line with 35-39 cycloid scales in series, and pelvic fin insertions slightly posterior to the dorsal fin insertion.

Distribution and Habitat: Native throughout most of North America from the Northern

Territories to Quebec in Canada, and in the United States from Montana to New Mexico in the west, east throughout the Missouri, Mississippi, St. Lawrence-Great Lakes, and

Ohio River drainages east of the Appalachian Mountains from Vermont to Texas. Widely distributed outside its native range due to popularity as a bait fish and the species tolerance and adaptability to a wide variety of habitats. Widely distributed throughout the

Dakotas. Well adapted to shallow pooled areas in wetland and stream habitats, but also occur in creeks, ponds and lakes.9,14 Tolerant of a wide variety of water quality variables including high levels of alkalinity, turbidity, ammonia and temperature, which are common characteristics of wetlands of the northern prairie region.2,12,14,17 An acclimation to higher water temperatures generally increases the upper lethal temperature, but a maximum lethal temperature has been reported as 34 °C (93.2 °F).2 Highly tolerant of hypoxic conditions, although not for prolonged periods of time.3,10 The species may also take advantage of differences in physiological tolerance to reduced dissolved oxygen concentrations to avoid a potential predator.1

Reproduction: Spawning takes place April to August, or when surface water temperatures range roughly 14.44-18.33 °C (58-65 °F).6,12 Spawning does not occur in water temperatures >30 °C (86 °F).2 Capable of reaching sexual maturity within 4-5 months of hatching under optimal conditions, but most do not spawn until age-1.2,11

Males and females may be of the same size when spawning, but females often spawn with larger males.11 Spawning generally begins before dawn and lasts until midmorning.6 334

Fractional spawners with time between spawning events ranging 2-16 days.6 Fecundity and the number of spawning events per female gradually decrease in water temperatures

>23.5 °C (74.3 °F).2 Six pairs of Fathead Minnows from Pennsylvania produced 16-26 clutches of eggs during the spawning season, with 9-1,136 eggs per clutch, to produce a total of 6,803-10,164 eggs per female per spawning season.6 The first clutch of eggs generally contains fewer eggs than later clutches.6 Nests are cavity-like and are constructed under rocks, logs, or other structures along the bottom substrate, with adhesive eggs deposited in one to two layers on the ceiling or underside of the structure.11

Eggs are roughly 1.15 mm (0.05 in) in diameter.11 Males provide parental care by actively guarding and cleaning the eggs until hatching.11,16 Males clean the eggs free of silt by rubbing them with their dorsal pad while also excreting a layer of mucus on them, which may increase egg survival.16 Egg production, quality and hatchability are known to be reduced in water with pH levels of 5.9 and lower.13 Hatching occurs within 4-5 days at

25 °C (77 °F).4

Age and Growth: Larvae approximately 4.75 mm (0.19 in) at hatching.11 Experience rapid growth, with juveniles reaching 45-50 mm (1.77-1.97 in) TL in 90 days.8,9 Mean lengths-at-age of individuals from North Dakota are reported as: age-1, 41 mm (1.61 in)

TL; age-2, 58 mm (2.28 in) TL.8 Males generally grow faster and attain greater lengths than females.8 Capable of reaching 101 mm (3.98 in) TL.12 Longevity 3 years, with majority of individuals only surviving until age-2.8

Food and Feeding: Opportunistic omnivore.8,9 Adult fathead minnows exhibit both particulate and filter feeding behaviors.7 Consumes a wide variety of prey items including detritus, zooplankton and macroinvertebrates.5,8,9,15 Detritus is an important prey item for 335 individuals of all sizes given that it allows individuals to survive when nutritional food sources are low.8,9,15 Juvenile and adult Fathead Minnows are known to select invertebrate prey over detritus when invertebrate prey is available and intra-specific competition is low.9

Literature Cited:

1. Abrahams, M.V., and J. Sloan. 2012. Risk of predation, variation in dissolved

oxygen, and their impact upon habitat selection decisions by Fathead Minnow.

Transactions of the American Fisheries Society 141:580-584.

2. Brungs, W.A. 1971. Chronic effects of elevated temperature on the fathead

minnow (Pimephales promelas Rafinesque). Transactions of the American

Fisheries Society 100:659-664.

3. Danylchuk, A.J., and W.M. Tonn. 2003. Natural disturbances and fish: local and

regional influences on winterkill of fathead minnows in boreal lakes. Transactions

of the American Fisheries Society 32:289-298.

4. Devlin, E.W., J.D. Brammer, R.L. Puyear, and J.M. McKim. 1996. Prehatching

development of the fathead minnow Pimephales promelas Rafinesque.

EPA/600/R-96/079, U.S. Environmental Protection Agency, Washington, D.C.

5. Duffy, W.G. 1998. Population dynamics, production, and prey consumption of

fathead minnows (Pimephales promelas) in prairie wetlands: a bioenergetics

approach. Canadian Journal of Fisheries and Aquatic Science 54:15-27.

6. Gale, W.F., and G.L. Buynak. 1982. Fecundity and spawning frequency of the

fathead minnow- a fractional spawner. Transactions of the American Fisheries

Society 111:35-40. 336

7. Hambright, K.D., and R.O. Hall. 1992. Differential zooplankton feeding

behaviors, selectivities, and community impacts of two planktivorous fishes.

Environmental Biology of Fishes 35:401-411.

8. Held, J.W., and J.J. Peterka. 1974. Age, growth, and food habits of the fathead

minnow, Pimephales promelas, in North Dakota saline lakes. Transactions of the

American Fisheries Society 103:743-757.

9. Herwig, B.R., and K.D. Zimmer. 2007. Population ecology and prey consumption

by fathead minnows in prairie wetlands: importance of detritus and larval fish.

Ecology of Freshwater Fish 16:282-294.

10. Klinger, S.A., J.J. Magnuson, and G.W. Gallepp. 1982. Survival mechanisms of

the central mudminnow (Umbra limi), fathead minnow (Pimephales promelas),

and Brook Stickleback (Culaea inconstans) for low oxygen in winter.

Environmental Biology of Fishes 7:113-120.

11. Markus, H.C. 1934. Life history of the blackhead minnow (Pimephales

promelas). Copeia 1934:116-122.

12. McCarraher D.B., and R. Thomas. 1968. Some ecological observation on the

fathead minnow, Pimephales promelas, in the alkaline waters of Nebraska.

Transactions of the American Fisheries Society 97:52-55.

13. Mount, D.I. 1973. Chronic effect of low pH on fathead minnow survival, growth

and reproduction. Water Research Pergamon Press 7:987-993.

14. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of

Conservation, Jefferson City. 337

15. Price, C.J., W.M. Tonn, and C.A. Paszkowski. 1991. Intraspecific patterns of

resource use by fathead minnows in a small boreal lake. Canadian Journal of

Zoology 69:2109-2115.

16. Smith, R.J. F., and B.D. Murphy. 1974. Functional morphology of the dorsal pad

in fathead minnows (Pimephales promelas Rafinesque). Transactions of the

American Fisheries Society 103:65-72.

17. Thurston, R.V., R.C. Russo, E.L. Meyn, R.K. Zajdel, and C.E. Smith. 1986.

Chronic toxicity of ammonia to fathead minnows. Transactions of the American

Fisheries Society 117:196-207.

338

Flathead Chub, Platygobio gracilis (Richardson, 1836)

Etymology and Synonyms: Platy = “flat”, gobio = name for the similar Eurasian cyprinid “gudgeon”; gracilis = slender, possibly referring to the species body shape.

Description: Body slender, elongate; slightly laterally compressed. Dorsally tan to light bronze; laterally silver with no distinct markings; ventrally silver to white; fins clear, transparent; lower lobe of caudal fin dusky gray with white edge. Head broad, short, dorsoventrally flattened; width greater than depth. Snout tapered to a point with a rounded tip protruding past upper jaw. Eye moderately small, placed laterally on head.

Mouth subterminal, large, slightly oblique, extending to anterior end of eye. Frenum absent. Barbels present, one on each corner of mouth; short, rounded. Lips thin.

Pharyngeal tooth pattern 2,4-4,2. Gill rakers short, 4-6. Dorsal fin with 8 rays, sickle shaped with pointed tip; anterior dorsal fin ray extends beyond last ray when depressed.

Adipose fin absent. Caudal peduncle slightly elongate, uniform in thickness. Caudal fin forked. Anal fin with 8 rays. Pelvic fins abdominal; insertions nearly equal to slightly posterior of dorsal fin insertion. Pectoral fins sickle shaped with pointed tip and 15-18 rays. Lateral line complete with 44-59 large cycloid scales in series. Spawning males may develop faint red coloration on fins, as well as small tubercles on dorsal side of head and body, the caudal peduncle and on all fins except the caudal fin.

Similar Species: Closely resembles the Plains Minnow and Western Silvery Minnow which lack barbels and sickle shaped dorsal and pectoral fins. Silver Chub have a larger eye and a rounded, short snout. Sicklefin Chub have a blunt snout and pectoral fins that extend beyond the insertion of the pelvic fins when depressed. Sturgeon Chub have a blunt snout, and fins with rounded tips lacking any sickle shape. 339

Distribution and Habitat: Widespread native distribution throughout four major North

American drainage systems: Mackenzie, Saskatchewan, Missouri-Mississippi, and the

Rio Grande.18 The species has declined throughout its historical range and within much of the lower Missouri River basin, likely due to barriers, impoundments, and channel modifications which reduce turbidity and alter the flow regime.6,12,13,14,23,24 Primarily occur in the main tributaries west of the Missouri River in the Dakotas, and rarely occur within the mainstem.1,12,14,22 Inhabits large turbid rivers with moderate to strong current over sand or silt substrates, which are common characteristics of Great Plains rivers and streams.13,19 Low velocity, shallow habitat is important for the Flathead Chub as it occurs frequently within channel border habitat at depths <1 m (3.28 ft) with current velocities

<0.25 m/s.24 The species is especially common in alkaline streams exhibiting large fluctuations in water levels and shifting sandy-loam substrates.18 Young are often found in large schools.20

Reproduction: Spawning in South Dakota likely takes place late June through early July, which is earlier than reported spawning seasons in Montana, Iowa and Kansas.5,9,12,17

Sexual maturity is reached between ages 2-4.8,13,17 Males from the upper Missouri River were all mature at 110 mm (4.33 in) TL at age-2, whereas females were not all mature until age-3 and 170 mm (6.69 in) TL.8 Broadcast, pelagic spawners.2 Spawning behavior consists of males swimming aggressively around females, and nudging them in the vent and abdominal areas.2 Information on where spawning takes place is scarce, but it is suggested to occur in pool habitat.17 Estimated fecundity of eight females from Montana ranged 360-735 eggs per female, however mature females from Iowa averaged 4,974 eggs/ovary.9,17 Mean number of eggs/female is known to increase with body length up to 340

130 mm (5.12 in) SL, and then slightly decline.17 Eggs orange in color, roughly 1.0-1.8 mm (0.04-0.07 in) in diameter, semi-buoyant, non-adhesive, and are carried with the current downstream during incubation and larval development.2,7,9,21 Hatching takes place within 6-7 days in a water temperature of 20-22 °C (68.0-71.6 °F).2

Age and Growth: Larvae roughly 5.5 mm (0.22 in) TL at hatching.11 Hatch date has been shown to be negatively correlated with growth rate, indicating that individuals who spawn later during the reproductive season exhibit slower growth rates than individuals who spawn earlier in the reproductive season.7 Mean length-at-age of individuals from the Moreau River in South Dakota are reported as: age-1, 60 mm (2.36 in) TL; age-2, 99 mm (3.90 in) TL; age-3, 133 mm (5.24 in) TL; age-4, 154 mm (6.06 in) TL; age-5, 169 mm (6.65 in) TL; age-6, 182 mm (7.17 in) TL.15 Individuals from the White River Basin in South Dakota indicated a more rapid increase in total length after age-4, which may possibly be due to changes in prey selectivity of large terrestrial insects.10 Capable of reaching 275 mm (10.83 in) TL in the Missouri River near the North Dakota-Montana border, and 370 mm (14.57 in) TL in Canada.8,12,16 Longevity 10 years.3

Food and Feeding: Prey consumption has been shown to be unaffected by increased levels of turbidity, likely due to morphological adaptations including barbels and a combination of visual and non-visual cues.4 Adult diets primarily consist of plant material and terrestrial insects that fall into and drift with the current.13 In high flow years, individuals <60 mm (2.36 in) TL were found to feed on Hemipterans and

Copepods, and predominantly on Coleopterans in years with average flow.8

Literature Cited: 341

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Bestgen, K.R., H.J. Crockett, M.R. Haworth, and R.M. Fitzpatrick. 2016.

Production of nonadhesive eggs by flathead chub and implications for

downstream transport and conservation. Journal of Fish and Wildlife Management

7:434-443.

3. Bishop, F.G. 1975. Observation on the fish fauna of the Peace River in Alberta.

The Canadian Field-Naturalist 89:423-430.

4. Bonner, T.H., and G.R. Wilde. 2002. Effects of turbidity on prey consumption by

prairie stream fishes. Transactions of the American Fisheries Society 131:1203-

1208.

5. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

6. Cross, F.B., R.L. Mayden, and J.D. Stewart. 1986. Fishes in the western

Mississippi drainage. Pages 363-412 in C.H. Hocutt and E.O. Wiley (editors). The

Zoogeography of North American Freshwater Fishes. John Wiley and Sons,

Toronto, Canada.

7. Durham, B.W., and G.R. Wilde. 2005. Relationship between hatch date and first-

summer growth of five species of prairie-stream cyprinids. Environmental

Biology of Fishes 72:45-54.

8. Fisher, S.J., D.W. Willis, M.M Olson, and S.C. Krentz. 2002. Flathead chubs,

Platygobio gracilis, in the upper Missouri River: the biology of a species at risk in

an endangered habitat. The Canadian Field-Naturalist 116:26-41. 342

9. Gould, W. 1985. Aspects of the biology of the flathead chub (Hybopsis gracilis)

in Montana. Great Basin Naturalist 45:332-336.

10. Harland, B.C. 2003. Survey of the fishes and habitat of western South Dakota

streams. M.S. Thesis, South Dakota State University, Brookings, South Dakota.

11. Haworth, M.R. 2015. Reproduction and recruitment dynamics of flathead chub

Platygobio gracilis relative to flow and temperature regimes in Fountain Creek,

Colorado. M.S. Thesis, Colorado State University, Fort Collins.

12. Hayer, C.A., N.L. Ahrens, and C.R. Berry, Jr. 2008. Biology of flathead chub,

Platygobio gracilis in three Great Plains rivers. Proceedings of the South Dakota

Academy of Science 87:185-196.

13. Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, 5.

Selected chubs and minnows (Cyprinidae): sicklefin chub (M. meeki), sturgeon

chub (M. gelida), silver chub (M. storeriana), speckled chub (M. aestivaus),

flathead chub (Platygobio gracilis), plains minnow (Hybognathus placitus), and

western silvery minnow (H. argyritis). Transactions of the Nebraska Academy of

Sciences and Affiliated Societies 21:99-108.

14. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry. 2007.

Zoogeographic patterns and faunal change of South Dakota fishes. North

American Naturalist 67:161-184.

15. Jones, S.J. 2018. Western prairie stream fisheries: an assessment of past and

present fish assemblage structure, biotic homogenization, and population

dynamics in western South Dakota streams. Unpublished M.S. Thesis, South

Dakota State University, Brookings, South Dakota. 343

16. Kristensen, J. 1980. Large flathead chub (Platygobio gracilis) from the Peace-

Athabasca Delta, Alberta, including a Canadian record. The Canadian Field

Naturalist 94:342.

17. Martyn, H.A., and J.C. Schmulbach. 1978. Bionomics of the flathead chub,

Hybopsis gracilis (Richardson). Proceedings of the Iowa Academy of Science

85:62-65.

18. Olund, L.J., and F.B. Cross. 1961. Geographic variation in the North American

cyprinid fish, Hybopsis gracilis. University of Kansas, Museum of Natural

History 13:323-348.

19. Peters, E.J., and R.S. Holland. 1994. Biological and economic analyses of the fish

communities in the Platte River: modifications and tests of habitat suitability

criteria for fishes in the Platte River. Completion Report. Federal Aid in Fish

Restoration Project Number F-78-R, Study III: Job III-2. University of Nebraska-

Lincoln.

20. Pflieger, W.L., and T.B. Grace. 1987. Changes in the fish fauna of the lower

Missouri River, 1940-1983. Pages 166-177 in Community and evolutionary

ecology of North American stream fishes. W. Matthews and D. Heins (editors).

University of Oklahoma Press, Norman.

21. Platania, S.P., and C.S. Altenbach. 1998. Reproductive strategies and egg types of

seven Rio Grande Basin cyprinids. Copeia 1998:559-569.

22. Quist, M.C, W.A. Hubert, and F.J. Rahel. 2004. Relations among habitat

characteristics, exotic species, and turbid-river cyprinids in the Missouri River 344

drainage of Wyoming. Transactions of the American Fisheries Society 133:727-

742.

23. Walters, D.M., R.E. Zuellig, H.J. Crockett, J.F. Bruce, P.M. Lukacs, and R.M.

Fitzpatrick. 2014. Barriers impede upstream spawning migration of flathead chub.

Transactions of the American Fisheries Society 143:17-25.

24. Welker, T.L., and D.L. Scarnecchia. 2004. Habitat use and population structure of

four native minnows (family Cyprinidae) in the upper Missouri and lower

Yellowstone rivers, North Dakota (USA). Ecology of Freshwater Fish 13:8-22.

345

Western Blacknose Dace, Rhinichthys obtusus (Agassiz, 1854)

Etymology and Synonyms: Rhinichthys = “snout fish”; obtusus = Latin for “blunt” and

“rostrum” in reference to the bluntly pointed snout. Rhinichthys atratulus (Hermann,

1804).

Description: Body fusiform, elongate, rather cylindrical anteriorly and slightly laterally compressed posteriorly. Dorsally blackish-brown to dark olive with small dark specks or mottling present; laterally gray to olive brown with small dark specks and mottling present, sometimes forming a diffuse dark lateral stripe extending through the eye and onto the snout; ventrally cream to silvery white; fins lightly pigmented with a light gray to cream tint. Head slightly dorsoventrally flattened. Snout elongate, bluntly pointed, slightly protrudes past mouth. Eye moderate, placed laterally on upper portion of head.

Mouth small, subterminal, slightly oblique; upper jaw scarcely exceeds lower jaw and extends to posterior end of nostril. Frenum present. Barbels sometimes present, small and inconspicuous; one at each corner of mouth. Lips fleshy, thick. Pharyngeal tooth pattern

2,4-4,2. Gill rakers short, 6-7. Dorsal fin with 8 rays. Adipose fin absent. Caudal peduncle elongate, uniform in thickness. Caudal fin moderately forked. Anal fin with 7 rays. Pelvic fins abdominal with 8 rays; insertions slightly anterior to insertion of dorsal fin. Pectoral fins with 13-15 rays. Lateral line complete with 60-75 small cycloid scales in series. Spawning males develop small tubercles on the lateral sides of the body, and larger tubercles on the head and pelvic fins. Males also develop nuptial coloration including a burnt orange lateral stripe. Juveniles similar to adults but with a more defined dark lateral stripe ending in a diffuse caudal spot. 346

Similar Species: Closely resembles the Longnose Dace. Longnose Dace have an upper jaw that greatly exceeds the lower jaw, and the distance from the tip of the snout to the anterior tip of the lower jaw is greater than or equal to the diameter of the eye.

Distribution and Habitat: A recent study looking at the mitochondrial DNA of

Blacknose Dace strongly suggests the designation of two different species: the Eastern

Blacknose Dace (Rhinichthys atratulus), and the Western Blacknose Dace (Rhinichthys obtusus) which occur within the Dakotas.4 Western Blacknose Dace are distributed primarily throughout the upper Mississippi and Ohio River drainages, as well as tributaries of the Great Lakes-St. Lawrence River (excluding the southeast of Lake

Ontario) and southeastern Atlantic drainages west of the Appalachians to approximately

Nebraska and the Dakotas.4 Adults prefer shallow, clear to slightly turbid waters of high gradient, small to medium headwater streams with moderate to rapid current and sand or gravel substrate.2,5 Fry often found within pool and shoal habitats with water velocities

<15 cm/sec over sand and silt substrate.3,11,12 Occasionally found in larger rivers and lakes.12 Overwinter in pool habitat.5 Upper lethal temperature reported as 29.3 °C (84.74

°F).12

Reproduction: Spawning takes place in the mornings from May to July, or when water temperatures hover 21 °C (69.8 °F).1,2,5,11 Spawning may also be dependent on photoperiod.8 Mature females and spawning males have been collected from swift flowing shallow waters roughly 76.2-152.4 mm (3-6 in) deep in areas with gravel substrate and overhanging vegetation or undercut stream banks.5,10 Sexual maturity occurs at age-2.2,5 Females inhabit deeper parts of pools until ready to spawn.7 Spawning behavior consists of two or more males courting a single female to the spawning 347 grounds.7 Males are not territorial towards females.6 The male wraps his caudal peduncle over the female, and the pair vibrates their bodies together for roughly two seconds at a time to initiate the release of eggs and milt.1,7 Females retreat back to deeper waters post spawning, and males continue to defend the spawning site.7 Fecundity increases with size of female.5 In Iowa, fecundity from females 38-61 mm (1.50-2.40 in) SL ranged 375-

2,500.5 Eggs from Iowa measured 1.2-1.5 mm (0.05-0.06 in) in diameter.5 Eggs from

Nebraska had mean diameters of 0.8 mm (0.03 in) in March and 1.3 mm (0.05 in) in

May.2

Age and Growth: Fry roughly 5 mm (0.20 in) TL at hatching.11,12 In Iowa, age-0 individuals reach an average of 24-27 mm (0.94-1.06 in) TL during their first year.5

Growth rates fastest May through July for age-1 and age-2 individuals.5 Ranges of standard lengths-at-age from Nebraska are reported as: age-1, 22-44 mm (0.87-1.73 in)

SL; age-2, 40-72 mm (1.57-2.83 in) SL; age-3, 68-81 mm (2.68-3.19 in) SL.2 Mature females attain greater mean standard lengths and weights than males.2 Capable of reaching 104 mm (4.09 in) TL.8 Longevity 3 years.5

Food and Feeding: Primarily carnivorous on benthic invertebrates.9 Consume large amounts of diptera larvae and adults, as well as smaller amounts of amphipod and isopod crustaceans, and ephemeroptera larvae.2,5,9 Young individuals feed in quiet, shallow waters over soft substrates, while adults often feed over riffles with some vegetation or within deep eddying pools.9 Majority of feeding takes place in the morning, with limited feeding activity at night.9

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 348

2. Bragg, R.J. 1978. The distribution and ecology of the Blacknose Dace, Rhinichthys atratulus

(Herman) in Nebraska. M.S. Thesis, University of Nebraska-Omaha, Omaha, Nebraska.

3. Gibbons, J.R.H., and J.H. Gee. 1972. Ecological segregation between Longnose and

Blacknose Dace (genus Rhinichthys) in the Mink River, Manitoba. Journal of the Fisheries

Research Board of Canada 29:1245-1252.

4. Kraczkowski, M.L., and B. Chernoff. 2014. Molecular phylogenetics of the Eastern and

Western Blacknose Dace, Rhinichthys atratulus and R. obtusus (Teleostei: Cyprinidae).

Copeia 2014:325-338.

5. Noble, R.L. 1965. Life history and ecology of Western Blacknose Dace, Boone

County, Iowa, 1963-1964. Iowa Academy of Science. 72:282-293.

6. Phillips, G.L. 1967. Sexual dimorphism in the Western Blacknose Dace, Rhinichthys

atratulus meleagris. Journal of the Minnesota Academy of Science 34:11-13.

7. Raney, E.C. 1940. Comparison of the breeding habits of two subspecies of black-nosed dace,

Rhinichthys atratulus (Hermann). The American Midland Naturalist 23:399-403.

8. Tarter, D.C. 1969. Some aspects of reproduction in the Western Blacknose Dace, Rhinichthys

atratulus meleagris Agassiz, in Doe Run, Meade County, Kentucky. Transactions of the

American Fisheries Society 98:454-459.

9. Tarter, D.C. 1970. Food and feeding habits of the Western Blacknose Dace, Rhinichthys

atratulus meleagris Agassiz, in Doe Run, Meade County, Kentucky. The American Midland

Naturalist 83:134-159.

10. Trautman, M.B. 1957. The fishes of Ohio. Ohio State University Press, Columbus, Ohio.

11. Traver, J.R. 1929. The habits of the Blacknosed Dace, Rhinichthys atronasus (Mitchell).

Journal of the Elisha Mitchell Scientific Society 45:101-129. 349

12. Trial, J.G., J.G. Stanley, M. Batcheller, G. Gebhart, O.E. Maughan, and P.C. Nelson. 1983.

Habitat suitability information: Blacknose Dace. U.S. Department of the Interior, Fish and

Wildlife Service. FWS/OBS-82/10.41.

350

Longnose Dace, Rhinichthys cataractae (Valenciennes, 1842)

Etymology and Synonyms: Rhinichthys = “snout fish”; cataractae = “of the cataract”, in reference to the type locality, Niagara Falls.

Description: Body fusiform, elongate, rather cylindrical anteriorly and slightly laterally compressed posteriorly. Dorsally dark brown to dark olive with mottling present; laterally olive to gray with dark mottled specks often forming a weak lateral stripe and a more defined nasal stripe anterior of eye in adults; ventrally cream to silvery white; fins lightly pigmented with slight cream to light golden-orange tint. Head slightly dorsoventrally flattened. Snout elongate, protrudes far beyond mouth. Eye moderate, placed laterally on upper portion of head. Mouth small, subterminal; upper jaw greatly exceeds lower jaw and extends to posterior end of nostril. Frenum present. Barbels sometimes present, small, and inconspicuous; one on each corner of mouth. Lips fleshy, thick. Pharyngeal tooth pattern 2,4-4,2. Gill rakers short, 6-10. Dorsal fin with 8 rays. Adipose fin absent.

Caudal peduncle elongate, uniform in thickness. Caudal fin moderately forked. Anal fin with 7 rays. Pelvic fins abdominal with 8 rays; insertions slightly anterior to insertion of dorsal fin. Axillary process sometimes present above origin of pelvic fins. Pectoral fins with 13-15 rays. Lateral line complete with 61-76 small cycloid scales in series.

Spawning males develop small tubercles on dorsal side of head and body, as well as larger tubercles on pelvic fin rays. Males also develop nuptial coloration, which includes orangish-red coloring on lips, cheeks, bases of all fins, and ventral side of body.

Spawning females also develop light orange to yellow nuptial coloration, but to a lesser extent than males. Juveniles similar in appearance to adults but with a dark spot present at base of caudal fin, and a more distinct, dark lateral stripe on body that fades with age. 351

Similar Species: Closely resembles the Western Blacknose Dace and Sturgeon Chub.

Western Blacknose Dace have an upper jaw that scarcely exceeds the lower jaw, as well as a snout that only slightly overhangs the mouth. The distance from the tip of the snout to the anterior tip of the lower jaw is less than the diameter of the eye. Sturgeon Chub lack a frenum and have a longitudinal ridge or keel present on the dorsal scales.

Distribution and Habitat: Native as far north as the Mackenzie River drainage near the

Arctic Circle to the northern Pacific Coast and Rocky Mountains into the Rio Grande drainage in northern Mexico, and east throughout the northern glaciated regions of the

Missouri, Mississippi, Great Lakes-St. Lawrence river drainages, and south throughout the Atlantic slope river drainages and Appalachian Mountains.2,9,16 Primarily distributed west of the Missouri River in the Dakotas, but also occurs in the northeast river drainages of North Dakota. Inhabits cold-water streams and northern lakes.10,19 Most abundant in steep gradient, cool headwaters or trout streams of large rivers.9,15 Primarily a riffle dwelling benthic species seeking areas with overhead cover.3,17 Generally avoids areas with little flow such as pools.4 Migrates to riffle habitats within six weeks after hatching where they remain for the duration of their lives.11,12,18 Both juveniles and adults have indicated preference for areas with higher water velocities and boulder substrate.12,18

Larger substrate such as large rocks and boulders are used as shelter from the fast current.8,17 Individuals known to select areas 10-19 cm (0.33-0.62 ft) in depth, with juveniles avoiding areas >20 cm (0.66 ft) deep, and adults avoiding areas <10 cm (0.33 ft) and >30 cm (0.98 ft) deep.18 Home ranges rather limited throughout the species lifetime, with individuals potentially only using 10-15 m (32.81-49.21 ft) of a stream.10,14 352

Fry seek out areas with overhead cover.3 Although rather intolerant of turbidity during long periods of time, Longnose Dace temporarily inhabit swift, turbid streams.9,16

Reproduction: Spawning takes place late spring to early summer or when water temperatures reach 11.7-19 °C (53.06-66.2 °F).5,6,9 Spawning in lotic habitats occurs over pits or riffles with loose, coarse substrate and water velocities of 45-60 cm/s near some form of vegetative cover.1,19 In lentic habitats, individuals spawn near inshore, wave- swept areas.5,9 Sexual maturity reached at age-2.9,17 Males actively defend territories prior to and during spawning by biting and butting any intruders.1 Males court females to the preferred spawning site by a combination of nudging and quivering movements.1

Fractional spawners; females capable of spawning more than six clutches of eggs per year.19 Fecundity positively correlated with size of female and somatic mass.19 A clutch of eggs is deposited within crevices in coarse substrate, and are likely guarded by one parent.17 Unfertilized eggs roughly 1.0-1.5 mm (0.04-0.06 in) in diameter, and increase to 2.1 mm (0.08 in) in diameter after water is absorbed during fertilization.7 Fertilized eggs transparent, demersal and adhesive.17 Eggs hatch within 3-4 days when incubated at a temperature of 21 °C (69.8 °F), and 7-10 days at 15.6 °C (60.08 °F).7,17

Age and Growth: Mean length at hatching ranges 4.5-5.9 mm (0.18-0.23 in) TL.7

Females generally live longer and reach greater lengths than males by age-3.13,15 Capable of reaching 178 mm (7.01 in) TL.4,10 Longevity 5 years.13,15

Food and Feeding: Opportunistic benthivore mainly consuming aquatic invertebrates and periphyton.9,13 Fry 0-49 mm (0-1.93 in) TL primarily consume algae, and feed on

Mayflies and Chironomids as they grow.9,13 Adults consume Ephemeropterans and

Dipterans such as Baetidae, Tendipedidae, Chironimidae, and Simulidae.9,13 Annelids, 353 crustaceans, mollusks and fish eggs have also been reported in diets, but to a lesser extent.4

Literature Cited:

1. Bartnik, V.G. 1970. Reproductive isolation between two sympatric dace,

Rhinichthys atratulus and Rhinichthys cataractae, in Manitoba. Journal of the

Fisheries Research Board of Canada 27:2125-2141.

2. Bartnik, V.G. 1972. Comparison of the breeding habits of two subspecies of

longnose dace, Rhinichthys cataractae. Canadian Journal of Zoology 50:83-86.

3. Bartnik, V.G. 1973. Behavioral ecology of the longnose dace, Rhinichthys

cataractae (Pisces, Cyprinidae), significance of the dace social organization.

Ph.D. Thesis, University of British Columbia.

4. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

5. Brazo, D.C., C.R. Liston, and R.C. Anderson. 1978. Life history of the longnose

dace, Rhinichthys cataractae, in the surge zone of eastern Lake Michigan near

Ludington, Michigan. Transactions of the American Fisheries Society 107:550-

556.

6. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

7. Cooper, J.E. 1980. Egg, larval and juvenile development of longnose dace,

Rhinichthys cataractae, and river chub, Nocomis micropogon, with notes on their

hybridization. Copeia 1980:469-478.

8. Culp, J.M. 1989. Nocturnally constrained foraging of a lotic minnow (Rhinichthys

cataractae). Canadian Journal of Zoology 67:2008-2012. 354

9. Edwards, E.A., H. Li, and C.B. Schreck. 1983. Habitat suitability index models:

Longnose dace. U.S. Department of the Interior, Fish and Wildlife Service.

FWS/OBS-82/10.33.

10. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

11. Gee, J.H., and K. Machniak. 1972. Ecological notes on a lake-dwelling population

of longnose dace (Rhinichthys cataractae). Journal of the Fisheries Research

Board of Canada 29:330-332.

12. Gee, J.H., and T.G. Northcote. 1963. Comparative ecology of two sympatric

species of dace (Rhinichthys) in the Fraser River system, British Columbia.

Journal of the Fisheries Research Board of Canada 20:105-118.

13. Gerald, J.W. 1966. Food habits of the longnose dace, Rhinichthys cataractae.

Copeia 1966:478-485.

14. Hill, J., and G.D. Grossman. 1987. Home range estimates for three North

American stream fishes. Copeia 1987:376-380.

15. Kuehn, J.H. 1949. A study of a population of longnose dace (Rhinichthys c.

cataractae). Minnesota Academy of Sciences 17:81-87.

16. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R.

Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina

Biological Survey Publication 1980-12.

17. McPhail, J.D., and C.C. Lindsey. 1970. Freshwater fishes of northwestern Canada

and Alaska. Bulletin of the Fisheries Research Board of Canada 173:1-381. 355

18. Mullen, D.M., and T.M. Burton. 1995. Size-related habitat use by longnose dace

(Rhinichthys cataractae). The American Midland Naturalist 133:177-183.

19. Roberts, J.H., and G.D. Grossman. 2001. Reproductive characteristics of female

longnose dace in the Coweeta Creek drainage, North Carolina, USA. Ecology of

Freshwater Fish 10:184-190.

356

Rudd, Scardinius erythrophthalmus (Linnaeus, 1758)

Etymology and Synonyms: Scardinius = Greek, referring to the Scardus mountain range between Moesia and Macedonia; erythrophthalmus = “red eye”, referring to the slight fleck of red coloration in the eye.

Description: Body deep, laterally compressed with arched back. Dorsally gold to brassy olive; laterally silver to gold with no distinct markings; ventrally silver to white; all fins with red to rosy-orange tint. Head small, triangular. Snout bluntly pointed. Eye moderately large, golden in color with a slight fleck of red coloration; placed laterally on head. Mouth small, terminal to superior; lower jaw slightly protruding beyond the upper jaw. Frenum absent. Barbels absent. Pharyngeal tooth pattern 3,5-5,3. Gill rakers 10-13 on first gill arch. Dorsal fin slightly falcate with 9-11 rays. Adipose fin absent. Caudal peduncle short, thick. Caudal fin forked with lower lobe slightly more elongate than upper lobe. Anal fin elongate, moderately falcate with 11-13 rays. Pelvic fins abdominal with 8-9 rays, insertions distinctly anterior to insertion of dorsal fin. Pectoral fins with

15-17 rays. Keel scaled, extending from pelvic fins to vent. Lateral line decurved and complete with 36-45 large cycloid scales rounded on the distal end in series. Spawning males develop bright red fins with distinct pale margins, and small tubercles on head, anterior end of body, and on the pectoral, dorsal and anal fin rays. Juveniles similar to adults, but with less coloration in the fins.

Similar Species: Closely resembles the Golden Shiner. Golden Shiner have a scaleless keel, faint yellow-orange to clear fins, and a dorsal fin with 7-9 rays. Common Shiner have a larger head, a large terminal and oblique mouth, a dorsal fin with 8 rays, a more slender and elongate caudal peduncle, and an anal fin with only 9 rays. 357

Distribution and Habitat: Native to the Europe, Asia Minor and interior Russia, with the first introduction into the United States occurring in 1916.2,3 In the early 1980’s, Rudd underwent an explosive anthropogenic dispersal within the United States, and were first identified in South Dakota during the early 1990’s.1,2 Not reported from North Dakota. In eastern South Dakota, Rudd have been collected from Interstate Lake, Lake Alice, Lake

Madison, East Vermillion Lake, and Mina Lake.1,6 In central South Dakota, they have been collected from Lake Francis Case, and from western South Dakota in Orman Dam,

Newell Lake, Sheridan Lake, Pactola Dam, Canyon Lake, and Angostora Reservior.1,6

Inhabits a variety of habitat types including rivers, streams, lakes and reservoirs in littoral areas with quiet, slow waters such as pools and backwaters with submerged aquatic vegetation.4,7,10 Little information is known about the habitat preferences of the species.

Tolerant to a wide variety of habitat types and variables, which likely contributes to its wide distribution across the United States.10

Reproduction: Spawning takes place during late spring to summer or when water temperatures exceed 15.55 °C (60 °F) in shallow vegetated waters.7 No nest is constructed, and no parental care is given. Fecundity of 86 females 138-1,701 g (0.30-

3.75 lbs.) from the Buffalo Harbor and the Niagara River near the northeastern end of

Lake Erie ranged 12,827-156,775 eggs.8 Eggs colorless to pale yellow, adhesive to vegetation, and roughly 0.43-1.23 mm (0.02-0.05 in) in diameter.4,7,9 Hatching occurs within 4-20 days depending on water temperature.7

Age and Growth: Mean lengths-at-age of individuals from Lake Alice in eastern South

Dakota are reported as: age-1, 64 mm (2.52 in) TL; age-2, 166 mm (6.54 in) TL; age-3,

258 mm (10.16 in) TL; age-4, 315 mm (12.40 in) TL; age-5, 354 mm (13.94 in) TL; age- 358

6, 373 mm (14.69 in) TL; age-7, 391 mm (15.39 in) TL; age-8, 405 mm (15.94 in) TL; age-9, 407 mm (16.02 in) TL; age-10, 421 mm (16.57 in) TL.1 Mean lengths-at-age of individuals from Newell Lake in western South Dakota are reported as: age-1, 57 mm

(2.24 in) TL; age-2, 128 mm (5.04 in) TL; age-3, 217 mm (8.54 in) TL; age-4, 278 mm

(10.94 in) TL; age-5, 306 mm (12.05 in) TL; age-6, 319 mm (12.56 in) TL; age-7, 325 mm (12.80 in) TL; age-8, 339 mm (13.35 in) TL.1 Growth of Rudd in South Dakota appears to be relatively fast compared to Rudd elsewhere, likely due to the high productivity of eutrophic waters and newly introduced populations that may still be expanding.1 Capable of reaching 617 mm (24.29 in) TL.11 Longevity 30 years.10

Food and Feeding: Individuals from the Buffalo Harbor and the Niagara River are considered primarily herbivorous, mainly consuming aquatic macrophytes during summer months, and supplementing with filamentous algae and fish, such as Emerald

Shiner in the spring and fall.8 Diets of Rudd from Zealand revealed that the species undergoes an ontogenetic diet shift, with young-of-the-year foraging on Cladocerans and

Chironomid larvae before switching to benthic invertebrates and some aquatic macrophytes as juveniles, and primarily macrophytes when they reached >150 mm (5.91 in) in length.5,8 Rudd >200 mm (7.87 in) FL had diets that consisted of >80% plant material.5 Feeding activity is significantly reduced during the spawning season, and is known to be positively related to water temperature.8

Literature Cited:

1. Blackwell, B.G., T.M. Kaufman, and W.H. Miller. 2009. Occurrence of rudd

(Scardinius erythrophthalmus) and dynamics of three populations in South

Dakota. Journal of Freshwater Ecology 24:285-291. 359

2. Burkhead, N.M., and J.D. Williams. 1991. An intergeneric hybrid of a native

minnow, the golden shiner, and an exotic minnow, the rudd. Transactions of the

American Fisheries Society 120:781-795.

3. Cahn, A.R. 1927. The European rudd (Scardinius) in Wisconsin. Copeia 162:5.

4. Haberlehner, E. 1988. Comparative analysis of feeding and schooling behaviour

of the Cyprinidae Alburnus alburnus, Rutilus rutilus and Scardinius

erythrophthalmus. International Review of Hydrobiology 73:537-546.

5. Hicks, B.J. 2003. Biology and potential impacts of rudd (Scardinius

erythrophthalmus L.) in New Zealand. Pages 49-58 in Managing Invasive

Freshwater Fish in New Zealand: proceedings of a workshop hosted by

Department of Conservation, Wellington, New Zealand.

6. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry, Jr. 2007.

Zoogeographic patterns and faunal change of South Dakota fishes. Western North

American Naturalist 67:161-184.

7. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources,

University of Nebraska-Lincoln.

8. Kapuscinski, K.L., J.M. Farrell, and M.A. Wilkinson. 2012. Feeding patterns and

population structure of an invasive cyprinid, the rudd Scardinius

erythrophthalmus (Cypriniformes, Cyprinidae), in Buffalo Harbor (Lake Erie) and

the upper Niagara River. Hydrobiologia 693:169-181.

9. Patimar, R., E Nadjafypour, M. Yaghouby, and M. Nadjafy. 2010. Reproduction

characteristics of a stunted population of rudd, Scardinius erythrophthalmus 360

(Linnaeus, 1758) living in the Anzali Lagoon (the southwest Caspian Sea, Iran).

Journal of Ichthyology 50:1060-1065.

10. Schofield, P.J., J.D. Williams, L.G. Nico, P. Fuller, and M.R. Thomas. 2005.

Foreign nonindigenous carps and minnows (Cyprinidae) in the United States- a

guide to their identification, distribution, and biology. U.S. Geological Survey

Scientific Investigations Report 2005-5041. 103p.

11. Šprem, N., D. Matulić, T. Treer, and I. Aničič. 2010. Short communication, a new

maximum length and weight for Scardinius erythrophthalmus. Journal of Applied

Ichthyology 26:618-619.

361

Creek Chub, Semotilus atromaculatus (Mitchill, 1818)

Etymology and Synonyms: Semotilus = “spotted banner”, in reference to the black spot present at the anterior dorsal fin base; atromaculatus = “black spot”.

Description: Body fusiform, robust, cylindrical anteriorly and laterally compressed posteriorly. Dorsally dark brown to olive; laterally bronze to silver with dark, diffuse lateral stripe ending in a spot at the base of the caudal fin; ventrally silver to white; dorsal fin with black spot present at the anterior base, remaining fins lightly pigmented. Head broad. Snout bluntly pointed. Eye large, placed laterally on upper portion of head. Mouth large, terminal; upper jaw extends past the anterior edge of the eye. Barbels present, small, flat and flap-like tucked into the fold between the upper lip and jaw. Frenum absent. Lips fleshy. Pharyngeal tooth pattern 2,5-4,2. Gill rakers short, 8. Dorsal fin with

8 rays, nearly straight to slightly convex distal end. Adipose fin absent. Caudal peduncle moderately thick. Caudal fin moderately forked. Anal fin with 8 rays. Pelvic fins abdominal with 8 rays, insertions anterior to insertion of dorsal fin. Pectoral fins with 16-

17 rays. Lateral line complete with 50-65 cycloid scales in series. Spawning males develop pink coloration on lower half of head, as well as reddish-orange coloring on the bases of the pectoral and pelvic fins. Males also display large tubercles present on the head and pectoral fins. Juveniles more silver in color with a more pronounced lateral stripe.

Similar Species: Easily distinguishable from other minnow species by the black spot on the anterior base of the dorsal fin present, one pair of flat, flap-like barbels in the fold between the upper lip and jaw, and a large terminal mouth. Hornyhead Chub have an upper jaw that extends to the anterior edge of the eye, and an anal fin with 7 rays. Lake 362

Chub have a relatively small mouth and lack a black spot on the anterior base of the dorsal fin.

Distribution and Habitat: Native to southern Manitoba and Quebec of Canada and much of the central and eastern United States from Montana and Wyoming in the west, east to the Atlantic Coast drainages, and south to the Gulf of Mexico.18 Widely distributed throughout the Dakotas in small streams and tributaries of the Missouri River basin. Inhabits small creeks and streams. Occasionally occur within ponds and lakes, although they are not considered preferred habitat.11 Prefer cool, clear to moderately turbid waters of high gradients in areas with gravel, cobble and rubble substrates, especially within well-defined pool-riffle areas with abundant cover.7,9,11 When exposed to higher turbidity, the use of overhead cover is known to decrease and activity levels are known to increase.9 Movement is also likely to decrease as habitat complexity and pool area is increased.20 Upper lethal temperature approximately 32 °C (89.6 °F), with a lower lethal temperature roughly 1.7 °C (35.06 °F).3,11 Juveniles known to be non-migratory, at least during their first year of life.10

Reproduction: Spawning takes place April through July, or when water temperatures reach 12.8 °C (55.04 °F).11,18 Spawning migrations upstream are generally less than 300 m (984.25 ft), but are capable of moving up to 1069.26 m (3508.07 ft).19,20 Sexual maturity begins entering their second year of life, or when individuals reach approximately 62.8 mm (2.47 in) TL.17 Males construct and defend a nest, or a redd, by pushing pieces of gravel or carrying them in its mouth to create a shallow depression.16

The nest may be of considerable size within gravel substrate often within shallow areas just above and below riffles.11,21 The spawning act consists of a single male embracing a 363 single female by wrapping his head and pectoral fins around her body, and the pair vibrating together to initiate the release of eggs and milt.16 After eggs settle into the nest, females will retreat back towards areas with cover or continue to spawn at another nest.16

Males guard the eggs until they have been completely covered and then abandon the nest.16 Fecundity generally increases with the size of the female.15,17 Fecundity in

Michigan ranged 3,500-5,000 eggs for females 127-152 mm (5.00-5.98 in) TL, while in

Ohio females 114-121 mm (4.49-4.76 in) TL ranged 4,193-4,671 mature ova.5,17,21

Mature eggs yellow-orange in color, and range 0.99-1.9 mm (0.04-0.07 in) in diameter with the size generally increasing with the size of the female.17 Hatching occurs in 10 days at 13 °C (55.4 °F).21

Age and Growth: Larvae roughly 6-7 mm (0.24-0.28 in) TL at hatching.2 Growth of age-0 individuals is rapid, highly variable and likely depends on food supply, stream flow, dissolved oxygen concentrations and the effects of pollution.7,10 Growth of age-0

Creek Chubs has also been shown to be related to the amount of total woody debris in a stream, likely because woody debris provides quality nutrient sources for aquatic invertebrates which make up a large portion of juvenile Creek Chub diets.7,13 Individuals from the Des Moines River in Iowa are reported as: age-1, 58 mm (2.28 in) TL; age-2, 95 mm (3.74 in) TL; age-3, 128 mm (5.04 in) TL.7 These lengths are slightly shorter than lengths recorded in northern Kansas.13 Capable of reaching 304.8 mm (12 in) TL.12

Longevity 8 years.12

Food and Feeding: Opportunistic, visual, generalist omnivores.8 Juveniles consume small terrestrial and aquatic insects during the day.1,6 Adult diets primarily consist of plant material, aquatic and terrestrial insects including Ephemeropterans and Coleopteran 364 adults, and in larger individuals, smaller fish.7 Adults are also known to consume

Annelids, crayfish, Gastropods and mussels.1,7,14 In intermittent environments, Creek

Chub have demonstrated the adaptive capability to increase their gut length to help them survive in in reduced resource environments without a cost to their condition.4

Literature Cited:

1. Barber, W.E., and W.L. Minckley. 1971. Summer foods of the cyprinid fish

Semotilus atromaculatus. Transactions of the American Fisheries Society

100:283-289.

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Brett, J.R. 1944. Some lethal temperature relations of Algonquin Park fishes.

University of Toronto Studies Biological Series, No. 52. The University of

Toronto Press, Ontario, Canada.

4. Christian, J.M., and G.L. Adams. 2014. Effects of pool isolation on trophic

ecology of fishes in a highland stream. Journal of Fish Biology 85:752-772.

5. Clark, C.F. 1943. Creek chub minnow propagation. Ohio Conservation Bulletin

7:12-13.

6. Copes, F. 1978. Ecology of the creek chub, Semotilus atromaculatus (Mitchill), in

northern waters. University of Wisconsin, Stevens Point, Museum of Natural

History, Fauna and Flora of Wisconsin Report Number 12:1-21.

7. Dinsmore, J.J. 1962. Life history of the creek chub, with emphasis on growth.

Iowa Academy of Science 69:296-301. 365

8. Evans, H.E. 1952. The correlation of brain pattern and feeding habits in four

species of cyprinid fishes. Journal of Comparative Neurology 97:133-142.

9. Gradall, K.S., and W.A. Swenson. 1982. Responses of brook trout and creek

chubs to turbidity. Transactions of the American Fisheries Society 111:392-395.

10. Katz, M., and W.C. Howard. 1955. The length and growth of 0-year class creek

chubs in relation to domestic pollution. Transactions of the American Fisheries

Society 84:228-238.

11. McMahon, T.E. 1982. Habitat suitability index models: creek chub. United States

Department of the Interior Fish and Wildlife Service. FWS/OBS-82/10.4.

12. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

13. Quist, M.C., and C.S. Guy. 2001. Growth and mortality of prairie stream fishes:

relations with fish community and instream habitat characteristics. Ecology of

Freshwater Fish 10:88-96.

14. Quist, M.C., M.R. Bower, and W.A. Hubert. 2006. Summer food habits and

trophic overlap of roundtail chub and creek chub in muddy creek, Wyoming. The

Southwestern Naturalist 51:22-27.

15. Reash, R.J., and T.M. Berra. 1986. Fecundity and trace-metal content of creek

chubs from a metal-contaminated stream. Transactions of the American Fisheries

Society 115:346-351.

16. Reighard, J. 1910. Methods of studying the habits of fishes, with an account of the

breeding habits of the horned dace. Bulletin of the US Bureau of Fisheries

28:1111-1136. 366

17. Schemske, D.W. 1974. Age, length and fecundity of the Creek Chub, Semotilus

atromaculatus (Mitchill), in central Illinois. The American Midland Naturalist

92:505-509.

18. Scott, W.B., and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Bulletin 184:507-510.

19. Storck, T., and W.T. Momot. 1981. Movements of the creek chub in a small Ohio

stream. Ohio Academy of Science 81:9-13.

20. Walker, R.H., and G.L. Adams. 2016. Ecological factors influencing movement

of creek chub in an intermittent stream of the Ozark Mountains, Arkansas.

Ecology of Freshwater Fish 25:190-202.

21. Washburn, G.N. 1945. Propagation of the creek chub in ponds with artificial

raceways. Transactions of the American Fisheries Society 75:336-350.

367

CHAPTER 11

FAMILY CATOSTOMIDAE

Introduction

The Sucker family, Catostomidae, are almost entirely native to North America. Of the 76 species within the family, only two species have distributions outside of North

America, the Asiatic Sucker, Myxocyprinus asiaticus, which is native to China, and the

Longnose Sucker, Catostomus catostomus, which is native to North America, but has expanded its distribution since the Pleistocene glacial period to eastern Siberia. Despite catostomids exhibiting a diverse array of body shapes, several species of suckers are often mistaken for larger, deeper bodied species of minnows from the family Cyprinidae, like the Common Carp, Cyprinus carpio. Although the head and body shapes may sometimes be relatively similar between the two families, catostomids are easily distinguishable from cyprinids by having a dorsal fin with ≥10 soft rays, a lack of barbels, and in most species of the family, a downward facing, “sucker-like” mouth.

Some species of sucker, like the critically endangered Razorback Sucker, Xyrauchen texanus, that is restricted to the Colorado River upstream of the Grand Canyon, exhibit distinct body morphology like a highly arched dorsal hump, which they have developed overtime in order to adapt to their preferred habitat of larger rivers with high water velocity.

The downward facing “sucker-like” mouth of catostomids is the main morphological feature that makes them immediately distinguishable from other families of fishes in the Dakotas. The lips tend to be enlarged, fleshy, covered with small taste 368 buds, and textured with either longitudinal grooves (plicate), numerous small bumps

(papillose), or have some sort of mixture of both. The variety of lip morphologies in suckers adds to their ability to seek out prey items and forage. Due to the position of the mouth, suckers have the ability to protrude their lips downward so that they can feed along the bottom substrate.

Throughout their distribution, catostomids inhabit a variety of habitat types including freshwater lakes, rivers and streams. In general, and in the Dakotas, species of catostomids that inhabit large river habitat in the display deep and more robust body types, while species that inhabit smaller streams often have a more streamlined and elongate body shape. The larger catostomids within the Dakotas such as the Quillback,

Carpiodes cyprinus, River Carpsucker, Carpiodes carpio, Highfin Carpsucker, Carpiodes velifer, Blue Sucker, Cycleptus elongatus, redhorse species (Moxostoma sp.) and buffalo fishes (Ictiobus sp.) are primarily found in larger rivers and lakes. The smaller and more streamlined species of suckers in the Dakotas such as Longnose Sucker, Catostomus catostomus, Mountain Sucker, Catostomus platyrhynchus, and White Sucker, Catosomus commersonii, primarily inhabit smaller rivers and streams.

The Dakotas have a great variety of catostomids that are widely distributed across both states. Five species of redhorse (Moxostoma sp.) primarily occur on the eastern side of the states with the exception of the Shorthead Redhorse, Moxostoma macrolepidotum, which is well-distributed east and west of the Missouri River. Bigmouth Buffalo, Ictiobus cyprinellus, Smallmouth Buffalo, Ictiobus bubalus, and the less common Black Buffalo,

Ictiobus miger, all occur within the Missouri River complex along with the Blue Sucker,

River Carpsucker and Highfin Carpsucker. Quillback also occur within the Missouri 369

River complex, but are also distributed within the Red River drainage. White Sucker are widespread across both North Dakota and South Dakota, whereas the Longnose Sucker has a disjunct distribution within the states, and occurs within the Missouri River in

North Dakota, and small areas of the northern Black Hills in South Dakota. The

Mountain Sucker is distinctly restricted to streams within the Black Hills of South

Dakota.

370

River Carpsucker, Carpiodes carpio (Rafinesque, 1820)

Etymology and Synonyms: Carpiodes = “carplike”; carpio = Latin name for “carp”, referring to the similarity of carp species.

Description: Body deep, laterally compressed with highly arched dorsal side. Dorsally olive to brown; laterally silver; ventrally cream to silvery white; dorsal and caudal fins light slate gray; ventral fins clear to light gray, with possible pink-orange pigmentation near bases. Head small. Subopercle with angled margin on lower corner. Snout short, bluntly rounded. Eye large, placed laterally on head. Mouth small, subterminal; upper jaw extends to or beyond anterior end of eye. Lips thin; lower lip forms an obtuse angle with a small, median nipple-like projection on tip. Barbels absent. Pharyngeal teeth numerous;

150-180 per side. Gill rakers long, fine, narrowly spaced. Dorsal fin elongate and falcate with 23-28 rays; anterior rays not greatly elongated with anterior serrated spine absent; anterior rays reach roughly 2/3 the length of the dorsal fin base when depressed. Adipose fin absent. Caudal peduncle short, thick. Caudal fin moderately forked. Anal fin with 7-8 rays. Pelvic fins abdominal with 8-10 rays; insertions posterior to insertion of dorsal fin.

Pectoral fins with 14-17 rays. Lateral line complete with 34-36 large cycloid scales in series. Spawning males develop small tubercles on dorsal and lateral sides of head, as well as on the pectoral and pelvic fin rays.

Similar Species: Closely resembles the Highfin Carpsucker. Highfin Carpsucker have anterior rays of the dorsal fin that are greatly elongated and extend to or nearly to the end of the dorsal fin base when depressed. Quillback display an upper jaw that does not extend to the anterior end of the eye, have a lower lip forming an acute angle, and lack a median nipple-like projection on the tip of the lower lip. Ictiobus species such as the 371

Bigmouth and Smallmouth Buffalo have a rounded margin on the lower corner of the subopercle.

Distribution and Habitat: Native throughout the Missouri, Mississippi and Ohio River basins from Montana in the west, east to western Pennsylvania, and south to Louisiana and New Mexico. Distributed throughout the Missouri River and its main tributaries in the Dakotas. The eight records from the Red River of the North may be misidentifications of Quillback, however these are unable to be verified as specimens were not vouchered from those collections. Primarily inhabits large to medium-sized rivers and reservoirs of low to moderate gradient in areas with quiet waters such as backwaters, pools, and runs with sand or silt substrate and woody debris.1,5,8

Reproduction: Spawning takes place mid-June to mid-July, or when water temperatures range from 19-24 °C (66.2-75.2 °F).10,18 Large numbers of individuals migrate upstream each spring from reservoirs into tributaries to spawn.4 Individuals are known to return to the same stream to spawn after being transplanted, highly suggesting spawning stream fidelity.4 Spawning grounds consist of shallow areas with little to moderate flow over sand and silt substrates.13 Spawning in South Dakota has been noted to be most successful during years when rising water levels during spring seep into flood marshes or low-lying meadows.17 Sexual maturity reached at age 4-8, and is related more to fish growth than age.8,18 Age at sexual maturity likely varies with latitude within the Missouri

River.6 Fractional spawners.2 No nest is constructed and no parental care is given.

Broadcast spawners.12 Fecundity of individuals from the Des Moines River in Iowa ranged 4,430 eggs from a 183 g (0.40 lb.) female, and 154,000 eggs from a 737 g (1.62 372 lb.) female.3 Eggs adhesive and roughly 1.2-1.5 mm (0.05-0.06 in) in diameter.1,12

Hatching occurs within 8-15 days.1

Age and Growth: Little difference in growth rates between sexes.18 Throughout the

Missouri River, mean lengths at ages 6-8 are known to increase with latitude.6 Growth rates during the first growing season are also positively correlated with latitude.6 Growth rates of age-1 individuals from the Des Moines River were faster when water levels were low.14 Growth of individuals inhabiting reservoirs is known to be greater than individuals from riverine habitat.16 Mean lengths-at-age from Lewis and Clark Reservoir in South

Dakota are reported as: age-1, 67 mm (2.64 in) TL; age-2, 130 mm (5.12 in) TL; age-3,

184 mm (7.24 in) TL; age-4, 226 mm (8.90 in) TL; age-5, 260 mm (10.24 in) TL; age-6,

286 mm (11.26 in) TL; age-7, 312 mm (12.28 in) TL.18 These growth rates are known to be slower than other Missouri River reservoirs.18 Minimum growth temperature 10 °C

(50 °F).10 Capable of reaching 496 mm (19.53 in) TL.18 Longevity 12 years, and increases from south to north in the Missouri River.6,18

Food and Feeding: Omnivorous facultative benthivore.11 Suction feeders, and selectively filter prey through their fine gill rakers and numerous pharyngeal teeth.9,11

Feeds on organic detritus, periphyton, zooplankton, phytoplankton, and aquatic insects associated with benthic habitats.7,15,18 Also known to consume Oligiochaetes and mollusks.15 Little variation in the diet by season, but has the ability to switch to lower quality prey items such as detritus when invertebrate abundance and availability is low.8,11,18

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 373

2. Behmer, D.J. 1967. Spawning periodicity of river carpsuckers, Carpiodes carpio.

Proceedings of the Iowa Academy of Science 72:252-262.

3. Behmer, D.J. 1969. A method of estimating fecundity; with data on river

carpsuckers, Carpiodes carpio. Transactions of the American Fisheries Society

98:523-524.

4. Bonneau, J.L., and D.L. Scarnecchia. 2002. Spawning-season homing of common

carp and river carpsucker. The Prairie Naturalist 34:13-20.

5. Braaten, P.J., and C.R. Berry, Jr. 1997. Fish associations with four habitat types in

a South Dakota prairie stream. Journal of Freshwater Ecology 12:477-489.

6. Braaten, P.J., and C.S. Guy. 2002. Life history attributes of fishes along the

latitudinal gradient of the Missouri River. Transactions of the American Fisheries

Society 131:931-945.

7. Brezner, J. 1958. Food habits of the northern river carpsucker. Progressive Fish-

Culturist 20:170-174.

8. Buchholz, M. 1957. Age and growth of river carpsucker in Des Moines River,

Iowa. Proceedings of the Iowa Academy of Science 64:589-600.

9. Eastman, J.T. 1977. The pharyngeal bones and teeth of catostomid fishes. The

American Midland Naturalist 97:68-88.

10. Fuiman, L.A. 1982. Family Catostomidae, suckers. Pages 345-435 in N.A. Auer,

editor. Identification of larval fishes of the Great Lakes basin with emphasis on

the Lake Michigan drainage. Great Lakes Fishery Commission, Special

Publication 82-3, Ann Arbor, Michigan. 374

11. Gido, K.B. 2001. Feeding ecology of three omnivorous fishes in Lake Texoma

(Oklahoma-Texas). The Southwestern Naturalist 46:23-33.

12. Jenkins, R.M. 1953. Growth histories of the principle fishes in Grand Lake (O’

The Cherokees), Oklahoma, through thirteen years of impoundment. Oklahoma

Fisheries Research Laboratory Report 34.

13. Jester, D.B. 1972. Life history, ecology and management of the river carpsucker,

Carpiodes carpio (Rafinesque), with reference to Elephant Butte Lake. University

of New Mexico Experiment Station Report 243:1-120.

14. Keeton, D. 1963. Growth of fishes in the Des Moines River, Iowa, with particular

reference to water levels. Ph.D. Dissertation, Iowa State University, Ames, Iowa.

15. Spiegel, J.R., M.C. Quist, and J.E. Morris. 2011. Trophic ecology and gill raker

morphology of seven catostomid species in Iowa rivers. Journal of Applied

Ichthyology 27:1159-1164.

16. Stucky, N.P., and H.E. Klassen. 1971. Growth and condition of the carp and the

river carpsucker in an altered environment in western Kansas. Transactions of the

American Fisheries Society 100:276-282.

17. Walburg, C.H. 1976. Changes in the fish populations of Lewis and Clark Lake,

1956-74, and their relation to water management and the environment. U.S.

Department of the Interior, Fish and Wildlife Service Research Report 79.

18. Walburg, C.H., and W.R. Nelson. 1966. Carp, river carpsucker, smallmouth

buffalo and bigmouth buffalo in Lewis and Clark Lake, Missouri River. U.S.

Department of the Interior, Fish and Wildlife Service, Research Report 69.

375

Quillback, Carpoides cyprinus (Lesueur, 1817)

Etymology and Synonyms: Carpiodes = carp-like; cyprinus = after the European island

Cyrus, where the carp was supposedly introduced; in reference to the species similarity to the carp.

Description: Body deep, laterally compressed with highly arched dorsal side. Dorsally silvery gray to light olive; laterally silver; ventrally cream to silvery white; dorsal and caudal fins light slate gray’ ventral fins light olive to gray with possible pink-orange pigmentation near bases. Head small. Snout short, blunt. Eyes large, placed laterally on head. Mouth small, subterminal; upper jaw barely extends to anterior end of eye. Lips thin; tip of lower lip lacking any nipple-like projection; lower lip forms an acute angle

(less than 90°). Barbels absent. Gill rakers long and thin. Dorsal fin elongate and falcate with 28-30 rays; anterior dorsal ray filamentous, falling between the middle end of the dorsal fin base, sometimes beyond, when depressed against the body; anterior serrated spine absent. Adipose fin absent. Caudal peduncle short, thick. Caudal fin forked. Anal fin with 7-8 rays. Pelvic fins abdominal with 9-10 rays; insertions posterior to insertion of dorsal fin. Lateral line complete with 36-40 large cycloid scales in series. Spawning males develop small tubercles on the ventral side of the head, lateral sides, and on the first dorsal fin ray. Larger spawning females occasionally develop tubercles on the lateral sides of the head.

Similar Species: Closely resembles the Highfin Carpsucker and River Carpsucker.

Highfin Carpsucker have anterior rays of the dorsal fin that are greatly elongated and reach to or nearly to the end of the dorsal fin base when depressed. Highfin Carpsucker also have a lower lip that forms an obtuse angle with a small, median nipple-like 376 projection present. River Carpsucker have anterior rays of the dorsal fin that extend about

2/3 the length of the dorsal fin base when depressed. River Carpsucker also have a lower lip that forms an obtuse angle with a small, median nipple-like projection present.

Distribution and Habitat: Native range spans across the Great Lakes and St. Lawrence

River, Hudson Bay, and Mississippi River basins from south-central Canada through the

Great Lakes, and over to the eastern Atlantic drainages, and south to the Gulf. Native to

South Dakota river drainages east of the Missouri River including the Red River of the

North, Upper Minnesota, Big Sioux, Vermillion, James, and Missouri basins, but have been documented in drainages west of the Missouri.6 Native to the Red River of the

North, Missouri, Sheyenne, and James river drainages in North Dakota.4,10,12 Rare in

Lake Sakakawea, and any other main stem Missouri River reservoirs.2,10 Inhabits clear to turbid quiet waters with little flow, and substrates of sand, gravel, silt, mud, or clay.1

Occurs in medium to low gradient large rivers, as well as lakes and creeks, sloughs, and sometimes small streams.1 Typically found in areas with little to no vegetation.8 The estimated mean critical thermal maximum for fish 45-47 mm (1.8-2.2 in) was 38.8°C

(102°F).8,9

Reproduction: Length of spawning period is thought to be prolonged, as tuberculate males have been reported as early as May and as late as September.1 Sexual maturity is known to occur at age 4-6 at a minimum of 313 mm (12.3 in) TL for males, and at age 6-

8 at a minimum of 386 mm (15.2 in) TL for females.11 Spawning is generally said to occur when water temperatures range from 19.0-28.0°C (66.2-82.4°F).1,16 In the early springtime, mature adults are known to migrate upstream into small creeks.1,15 Individuals are known to be monospawners per spawning season.2,14 Fecundity and egg size known to 377 increase with size of the female, with females capable of producing 15,235-63,779 eggs.8,11,16 Eggs are deposited over substrates of sand, gravel, and mud in quiet waters, with no nest prepared or parental care provided.1 Hatching occurs with 11-18 days in waters 16°C (61°F).5,8,11

Age and Growth: Males generally larger than females at each age.1,16 Average length is

356 mm (14 in) TL, but the species is capable of reaching up to 610 mm (24 in) TL.1,7

Mean lengths-at-age from Ohio are reported as: age-1, 84 mm (3.31 in) TL; age-2, 156 mm (6.14 in) TL; age-3, 206 mm (8.11 in) TL; age-4, 242 mm (9.53in) TL; age-5, 280 mm (11.02 in) TL; age-6, 312 mm (12.28 in) TL; age-7, 342 mm (13.46 in) TL; age-8,

368 mm (14.49 in) TL; age-9, 398 mm (15.67 in) TL; age-10, 428 mm (16.85 in) TL; age-11, 448 mm (17.64 in) TL.16 Long-lived species; longevity 11 years.16

Food and Feeding: Quillbacks are generalist, scavenger, benthic feeders and forage freely on plant materials, insect larvae such as chironomid larvae, and a variety of snails, clams, worms, and zooplankton.1,3,10,14 It has been suggested that Quillback compete with

Channel Catfish for invertebrate prey items.10,13

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Benson, N.G. 1968. Review of fishery studies on Missouri River main stem

reservoirs. U.S. Department of the Interior Fish and Wildlife Service, Bur. Of

Sport Fisheries and Wildlife, Res. Rept. 71 iv +61p.

3. Cahn, A.R. 1927. An ecological study of the southern Wisconsin fishes. The

brook silverside (Labidesthes sicculus) and the cisco (Leucichthys artedi) in their

relations to the region. Ill. Boil. Monogr. 11(1):1-151. 378

4. Copes, F.A. 1965. Fishes of the Red River tributaries of North Dakota. M.S.

Thesis, University of North Dakota. 65p.

5. Curry, K.D., and A. Spacie. 1984. Differential use of stream habitat by spawning

catostomids. American Midland Naturalist 111:267-269.

6. Hoagstrom, C.W. 2006. Zoogeographic patterns and faunal change of South

Dakota fishes. PhD. Dissertation. South Dakota State University, Brookings, SD.

7. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

8. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

9. Mundahl, N.D. 1990. Heat death of fish in shrinking stream pools. American

Midland Naturalist.

10. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

11. Parker, B.R., and W.G. Franzin. 1991. Reproductive biology of the quillback,

Carpiodes cyprinus, in a small prairie river. Canadian Journal of Zoology

69:2133-2139.

12. Peterka, J.J. 1978. Fishes and fisheries of the Sheyenne River, North Dakota.

Proceedings of the North Dakota Academy of Science 32 (Part II):29-44.

13. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p. 379

14. Spiegel, J.R., M.C. Quist, and J.E. Morris. 2011. Trophic ecology and gill raker

morphology of seven catostomid species in Iowa rivers. Journal of Applied

Ichthyology 27:1159-1164.

15. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press. 683 p.

16. Woodward, R,L., and T.E. Wissing. 1976. Age, growth, and fecundity of the

Quillback (Carpiodes cyprinus) and Highfin (C. velifer) Carpsuckers in an Ohio

stream. Transactions of the American Fisheries Society 105:411-415.

380

Highfin Carpsucker, Carpiodes velifer (Rafinesque, 1820)

Etymology and Synonyms: Carpiodes = “carplike”; velifer = “sailbearer”, in reference to the elongate and filamentous anterior dorsal fin rays that appear as a sail when near the surface of the water.

Description: Body deep, laterally compressed with highly arched dorsal side. Dorsally olive to brown; laterally silver; ventrally cream to silvery white; fins generally clear but may display a slate gray tint. Head small. Subopercle with angled margin on lower corner. Snout short, bluntly rounded. Eye large, placed laterally on head. Mouth small, subterminal; upper jaw extends to or beyond anterior end of eye. Lips thin; lower lip forms an obtuse angle with a small, median nipple-like projection on tip. Barbels absent.

Gill rakers long. Dorsal fin elongate and falcate with 22-27 rays; anterior rays greatly elongated with anterior serrated spine absent; anterior rays reach to or nearly to end of dorsal fin base when depressed. Adipose fin absent. Caudal peduncle short, thick. Caudal fin moderately forked. Anal fin with 8-9 rays. Pelvic fins abdominal with 9-10 rays; insertions posterior to insertion of dorsal fin. Pectoral fins with 14-17 rays. Lateral line complete with 33-35 large cycloid scales in series. Spawning males develop small tubercles on dorsal and lateral sides of head, snout, and majority of body, as well as on the pectoral and pelvic fin rays.

Similar Species: Closely resembles the River Carpsucker. River Carpsucker have anterior rays of the dorsal fin that are not greatly elongated, and only extend roughly 2/3 the length of the dorsal fin base when depressed. Quillback also have anterior rays of the dorsal fin that are shorter than the dorsal fin base when depressed, display an upper jaw that does not extend to the anterior end of the eye, have a lower lip forming an acute 381 angle, and lack a median nipple-like projection on the tip of the lower lip. Ictiobus species such as the Bigmouth and Smallmouth Buffalo, have a rounded margin on the lower corner of the subopercle.

Distribution and Habitat: Native throughout central North America in the lower

Missouri, Mississippi and Ohio River basins from Minnesota in the north and south to the

Gulf of Mexico. Has declined throughout much of its range, especially from the western edge of the species historical distribution.1,3,6 Range reduction is known to be associated with water pollution, siltation, and migratory barriers such as dams, which likely explains the declining and restricted range of the species in South Dakota.6,13 In South Dakota, the species is now known to be absent from the Big Sioux River drainage where it was once present.6,8 More common throughout the lower Missouri River basin than in the recreational river sections of South Dakota and Nebraska.2 Absent from North Dakota.

Riverine specialist.9 Inhabits medium to large-sized rivers and streams of swift velocities, with a preference for moderately deep to deep pools or areas adjacent to river channels.1,3,9,16 Prefers waters of relatively low turbidity over sand and silt substrates with little to no aquatic vegetation.1,10,11 When water levels are low, the species may generally be found in riffle areas.1 Less tolerant of high turbidity and siltation than other carpsuckers.1,11

Reproduction: Information on spawning behavior and reproductive habits is scarce.

Spawning takes place early spring, typically during May, when large numbers of individuals begin to migrate to shallow and overflow areas of streams and ponds to spawn.1,5 Spawning also known to occur over shallow riffles with gravel substrates.11 In

Ohio, sexual maturity occurs when individuals reach roughly 229 mm (9.02 in) TL or 382 less.16 In Iowa, sexual maturity is said to be reached during the third year of life.5

Fecundity of four females age 5-8 ranged 41,644 eggs from a 275 mm (10.83 in) TL, 247 g (0.54 lb.) female, and 62,355 eggs from a 300 mm (11.81 in) TL, 312 g (0.69 lb.) female.18 Eggs from Wisconsin averaged 1.2 mm (0.05 in) in diameter.1

Age and Growth: Smallest of the carpsuckers.4 Greatest amount of growth occurs during first year of life.17 Males typically attain greater lengths than females at each annulus.18

Mean lengths-at-age from the Des Moines River in Iowa are reported as: age-1, 88.9 mm

(3.5 in) TL; age-2, 132.08 mm (5.2 in) TL; age-3, 165.1 mm (6.5 in) TL; age-4, 195.58 mm (7.7 in) TL; age-5, 220.98 mm (8.7 in) TL; age-6, 259.08 mm (10.2 in) TL; age-7,

279.4 mm (11.0 in) TL; age-8, 304.8 mm (12.0 in) TL.17 Highfin Carpsuckers from Ohio and the Illinois River in Oklahoma had higher growth rates than individuals from the Des

Moines River, likely due to longer growing seasons.7,17,18 Capable of reaching 423 mm

(16.65 in) TL.14,15 Longevity 9 years.12

Food and Feeding: Benthic omnivore. Have more specialized diets compared to the

River Carpsucker and Quillback, consuming a fewer variety of prey items.15 Diet dominated by detritus, algae and aquatic invertebrates associated with benthic habitats such as Chironomid larvae, Oligiochaetes and mollusks.15

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin

2. Berry, Jr., C.B., and B. Young. 2004. Fishes of the Missouri national recreational

river, South Dakota and Nebraska. Great Plains Research 14:89-114.

3. Cross, F.B. 1967. Handbook of fishes of Kansas. Miscellaneous Publication No.

45. Museum of Natural History, University of Kansas, Lawrence. 383

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

5. Harlan, J.R., and E.B. Speaker. 1951. Iowa fish and fishing. State Conservation

Commission, Des Moines, Iowa.

6. Hoagstrom, C.W., C.A. Hayer, J.G. Kral, S.S. Wall, and C.R. Berry Jr. 2006. Rare

and declining fishes of South Dakota: a river drainage scale perspective.

Proceedings of the South Dakota Academy of Science 85:171-211.

7. Jenkins, R.M., E.M. Leonard, and G.E. Hall. 1952. An investigation of the fishery

resources of the Illinois river and pre-impoundment study of Tenkiller Reservoir,

Oklahoma. Oklahoma Fisheries Research Laboratory Report No. 26.

8. Lee, D.S., and S.P. Platania. 1978. Carpiodes velifer (Rafinesque), highfin

carpsucker. In D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister,

and J.R. Stauffer, Jr. Atlas of North American Freshwater Fishes. North Carolina

State Museum of Natural History, Raleigh.

9. Lyons, J. 1996. Effects of flow regulation and restriction of passage due to

hydroelectric project operation on the structure of fish and invertebrate

communities in Wisconsin’s large river systems. 1996 Progress Report, Phase 1.2:

Development of an Index of Biotic Integrity. Department of Natural Resources,

Madison, Wisconsin.

10. Miller, R.J., and H.W. Robinson. 2004. Fishes of Oklahoma, revised edition.

University of Oklahoma Press, Norman.

11. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 384

12. Quist, M.C., and J.R. Spiegel. 2012. Population demographics of catostomids in

large river ecosystems: effects of discharge and temperature on recruitment

dynamics and growth. River Research and Applications 28:1567-1586.

13. Smith, P.W. 1979. The fishes of Illinois. University of Illinois Press, Urbana.

14. Spiegel, J.R., M.C. Quist, and J.E. Morris. 2010. Estimating age of highfin

carpsucker, quillback carpsucker, and river carpsucker. Journal of Freshwater

Ecology 25:271-278.

15. Spiegel, J.R., M.C. Quist, and J.E. Morris. 2011. Trophic ecology and gill raker

morphology of seven catostomid species in Iowa rivers. Journal of Applied

Ichthyology 27:1159-1164.

16. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

17. Vanicek, D. 1961. Life history of the quillback and highfin carpsuckers in the Des

Moines River. Proceedings of the Iowa Academy of Science 68:238-246.

18. Woodward, R.L., and T.E. Wissing. 1976. Age, growth, and fecundity of the

quillback (Carpiodes cyprinus) and highfin (C. velifer) carpsuckers in an Ohio

stream. Transactions of the American Fisheries Society 105:411-415.

385

Longnose Sucker, Catostomus catostomus (Forster, 1773)

Etymology and Synonyms: Catostomus = “inferior mouth”; catostomus = “inferior mouth”.

Description: Body fusiform, elongate, slightly dorso-ventrally flattened. Dorsally dark gray to black-olive; laterally gray to brown abruptly changing to silvery-white ventrally; dorsal and caudal fin dark gray, other fins light gray to clear. Head long, slender, flattened between eyes. Snout long, pointed with blunt tip; extends well beyond upper lip.

Eye small, positioned high on posterior end of head. Mouth inferior, large. Lips fleshy, heavily papillose; lower lip completely divided by ventral notch forming an acute angle.

Pharyngeal teeth comb-like, 44-55; located in a single row on the fifth gill arch with.13

Gill rakers short; anterior row of first arch with 22-27 rakers, including rudimentary ones.13 Dorsal fin slightly concave with 9-12 soft rays. Adipose fin absent. Caudal peduncle short. Caudal fin moderately forked. Anal fin with 7 rays. Pelvic fin with 9 rays; insertion posterior to anterior end of dorsal fin. Pectoral fins rounded. Lateral line complete with 95-115 cycloid scales crowded anteriorly. Spawning males develop a dark red lateral stripe and tubercles on the head, anal fin, and caudal fin. Juveniles similar to adults, occasionally with three large black lateral blotches.

Similar Species: Closely resembles the Mountain and White Sucker. Mountain Sucker have a lateral line with 75-92 cycloid scales, inner cartilaginous ridges present on both jaws, lateral notches present on each corner of the mouth, and the lower lip lobe is incompletely divided. White Sucker with a slightly pointed snout barely extending past the upper lip, and 55-75 cycloid scales in lateral line series. 386

Distribution and Habitat: Native throughout Alaska and Canada and from the northwestern United States from Washington in the west, east to the Missouri River in the Dakotas, and south to Colorado. Also native to the Great Lakes region and New

England area. Longnose Sucker are the only species of North American sucker that is also native to northern Asia. Occurs throughout the Missouri River in North Dakota, and is restricted to small areas of the northern Black Hills in the Belle Fourche river drainage in South Dakota.1 Occupy a wide range of habitats, including small to medium sized rivers and streams, as well as lakes and reservoirs. Benthic dweller. Prefers cool, clear water with little to no turbidity and sand or gravel substrates. Most common at depths near 30-40 m (98-131 ft) in oligotrophic lakes and reservoirs, but have been found at depths up to 183 m (600 ft) in Lake Superior.8,10,12 Optimal water temperature for adults is 10-15°C (50-59°F).5,8 Most larvae 11-18 mm (0.43-0.71 in) TL remain in substrate near the spawning grounds for 9-14 days before moving into deeper water.4,6,12 Juveniles often found schooling in shallow backwater or lentic areas with dense aquatic vegetation.11

Reproduction: Spawning migrations and behavior initiated by water temperature and discharge.2,6 Migrate upstream or into smaller tributaries to spawn when water temperatures near 5-9°C (41-48°F).12 Sexual maturity is reached by age-6. Males reach sexual maturity at roughly age-4, and females age-5.4,9 Spawning takes place during the day in slow moving water or pools just downstream of riffles over gravel substrate when water temperatures reach 10-15°C (50-59°F).6,11,12 No nest is built and no parental care is given. Spawn in groups up to ten, with usually one larger female instigating a chase from several smaller males.7,11 Once positioned still over the spawning grounds, males nudge 387 females with their snout and vibrate their bodies to instigate egg release. Eggs are broadcast, light yellow in color, roughly 2.2 mm (0.09 in) in diameter, demersal, and adhesive to the gravel substrate.3,7 Fecundity dependent on the size of the female. Eggs hatch within 10-20 days. At a mean temperature of 12.2°C, embryos hatched after 14 days.12

Age and Growth: Females typically larger, grow more slowly, and live longer than males. Growth varies greatly after sexual maturity is reached.11 Mean lengths at age for both sexes combined in Yellowstone Lake, Wyoming were recorded as: age-1, 41 mm

(1.61 in) TL; age-2, 83 mm (3.27 in) TL; age-3, 147 mm (5.79 in) TL; age-4, 208 mm

(8.19 in) TL; age-5, 267 mm (10.51 in) TL; age-6, 313 mm (12.32 in) TL; age-7, 347 mm

(13.66 in) TL; age-8, 370 mm (14.57 in) TL.11 Average length 254-406 mm (10-18 in)

TL. Capable of reaching lengths of 609 mm (24 in) TL. Long lived species. Longevity is

8-11 years.2,12

Food and Feeding: Specialized, pelagic and benthic omnivores. Longnose Sucker feed more frequently in the pelagic zone than other species of suckers in the Dakotas. Fry feed on zooplankton and diatoms. Diet from juvenile to adult stages does not change drastically.11 Adults primarily consume algae and detritus, and often migrate near shorelines during the night to feed. Adults also known to consume amphipods, various forms of small aquatic insects, and zooplankton such as cladocerans, isopods, and copepods. Juveniles primarily feed on amphipods, zooplankton, and Tendipedidae larvae.11

Literature Cited: 388

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Barton, B.A. 1980. Spawning migrations, age, and growth, and summer feeding

of white and longnose suckers in an irrigation reservoir. Canadian Field Naturalist

94(3):300-304.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

5. Brown, C.J.D., and R.J. Graham. 1953. Observations on the Longnose Sucker in

Yellowstone Lake. Transactions of the American Fisheries Society 83(1):38-46.

6. Childress, E.S., R. Papke, and P.B. McIntyre. 2016. Spawning success and early

life history of longnose suckers in Great Lakes tributaries. Ecology of Freshwater

Fish 25:393-404.

7. Dion, R., M. Richardson, L. Roy, and F.G. Whoriskey. 1993. Spawning patterns

and interspecific matings of sympatric white (Catostomus commersoni) and

longnose (C.catostomus) suckers from the Gouin reservoir system, Quebec.

Canadian Journal of Zoology 72:195-200.

8. Edwards, E.A. 1983. Habitat suitablility index models: Longnose sucker. U.S.

Department of Interior, Fish and Wildlife Servive. FWS/OBS-82/10.35. 21pp.

9. Rawson, D.S., and C.A. Elsey. 1950. Reduction in the longnose sucker population

of Pyramid Lake, Alberta, in an attempt to improve angling. Transactions of the

American Fisheries Society 78(1):13-31. 389

10. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

11. Swanson, R.D. 1981. Some aspects of the biology of the longnose sucker in

Yellowstone Lake, Yellowstone National Park, Wyoming. Master’s thesis,

University of Wyoming, Laramie, Wyoming.

12. Walton, B.D. 1980. The reproductive biology, early life history, and growth of

white suckers, Catostomus commersoni, and longnose suckers, C. catostomus, in

the Willow Creek-Chain Lakes System, Alberta. Master’s thesis, University of

Alberta, Edmonton, Alberta.

13. Weisel, G.F. 1962. Comparative study of the digestive tract of a sucker,

Catostomus catostomus, and a predaceous minnow, Ptychocheilus oregonense.

The American Midland Naturalist, 68(2):334-346.

390

White Sucker, Catostomus commersonii (Lacepède, 1803)

Etymology and Synonyms: Catostomus = inferior mouth; commersonii = referring to

Dr. Philbert Commerson, a French naturalist.

Description: Body fusiform, elongate, slightly oval in cross section. Dorsal and lateral sides dark gray, brown, or olive with varying small, dark mottling; ventrally silver-white; dorsal and caudal fin light gray; anal, pelvic and pectoral fins clear with faint orange coloring. Head conical, slightly dorso-ventrally flattened and convex between eyes. Snout blunt, barely extending past upper lip. Eye small. Mouth inferior, large. Lips fleshy, heavily papillose; lateral notch absent. Lower lip almost twice as thick as upper lip, and forms an acute angle. Pharyngeal teeth thin, roughly 55-58 per arch. Dorsal fin nearly straight with 10-13 rays; height nearly equal to base length. Adipose fin absent. Caudal peduncle short and thick. Caudal fin forked. Anal fin with 7 rays. Pelvic fin with 10-11 rays; insertion posterior to anterior end of dorsal fin. Pectoral fins rounded with 16-19 rays. Lateral line complete with 55-75 cycloid scales. Scales smaller and more crowded anteriorly on body. Spawning males develop large tubercles on anal and caudal fin, and smaller tubercles on head and anterior end of body. Males also develop a thick, dark lateral stripe. Juveniles similar to adults, but with 3-4 large blotches on lateral sides.

Similar Species: Closely resembles the Mountain and Longnose Sucker. Mountain

Sucker have a lateral line with 75-92 cycloid scales, inner cartilaginous ridges present on both jaws, lateral notches present on each corner of the mouth, and a lower lip lobe that is completely divided. Longnose Sucker have a longer snout extending well beyond the upper lip and 95-115 cycloid scales in the lateral line. Redhorse species (Moxostoma spp.) in the Dakotas also resemble the White Sucker, however they have larger scales in 391 their lateral lines (<50) and more plicate lips. The Northern Hogsucker is brown to olive in color and heavily mottled with 4-6 dark irregular saddles.

Distribution and Habitat: One of the most widespread, adaptable, and abundant species of freshwater fish throughout North America.10 Native to much of Canada and the United

States east of the Rocky Mountains, from Montana in the northwest, east to the Atlantic

Coast, and south to northern Alabama and New Mexico.7 Widespread in both North and

South Dakota. Inhabits a wide variety of habitats including lakes, reservoirs, and small to medium sized rivers and streams. Adults prefer pool habitat in low to moderate gradient rivers and streams, and areas with slow to moderate velocity (~40 cm/sec) in lakes and reservoirs with clear water and gravel, sand, or silt substrates.11 Often associated with vegetative cover or woody debris. Able to tolerate wide ranges of turbidity and siltation.11

In the James and Missouri Rivers in North Dakota, juveniles and adults have been reported in turbidities of 50-135 JTU’s.11 Optimum water temperature varies geographically, however their wide distribution suggests that they can survive water temperatures as low as 1-2°C (33.8-35.6°F), and have a critical thermal maximum of

31.6°C (88.88°F).9,11 Juveniles occupy shallow backwaters or riffle areas with moderate velocity.11 Although the species migrate to spawn, there is strong evidence that the species remain in small home ranges for most of the year.5

Reproduction: Spawning migrations begin when water temperatures reach 7-10°C

(44.6-50°F) and usually take place at night. The species may migrate up to 40 km (25 mi) to spawn in tributaries or shallow areas near riffles with moderate water velocities over sand or gravel substrate.5 Sexual maturity is reached at ages 2-4, but depends upon location and availability of food.3,4,11 Males mature 1-2 years faster than females.10,3 392

Spawning takes place in spring typically from April-May. A single female is often wedged between two males on each side until eggs are released and fertilized.8 Spawning activity known to increase at dusk and dawn. No nest is prepared and no parental care is given. Eggs demersal, adhesive, and roughly 2-3 mm (0.07-0.12 in) in diameter.3

Fecundity from females 406-533 mm (16-21 in) TL ranged 20,000-50,000 eggs.3

Maximum hatching success has been reported at 15°C (59°F), with success decreasing in temperatures <9°C (48.2°F) or >17°C (62.6°F).6,11 Spawning may occur multiple times within one season, but may occur annually if environmental conditions are not adequate.5

Age and Growth: Females typically larger than males. Growth varies greatly among populations. Average length 203-305 mm (8-12 in) TL, but capable of reaching 610 mm

(24 in) TL. Longevity 10-17 years.2

Food and Feeding: Benthic feeder. Most feeding occurs at night when the species migrates near shore. Have the ability to sort and filter out food within the buccal cavity.1

Detritus is an important part of the juvenile diet, along with aquatic invertebrates and microcrustaceans such as cladocerans.1 Adults consume a wide variety of prey items including plankton, aquatic benthic invertebrates, vegetation, small fish, and fish eggs.

Literature Cited:

1. Ahlgren, M.O. 1990. Diet selection and the contribution of detritus to the diet of

the juvenile White Sucker (Catostomus commersoni). Canadian Journal of

Fisheries and Aquatic Sciences 47:41-48.

2. Beamish, R.J. 1973. Determination of age and growth of populations of the White

Sucker (Catostomus commersoni) exhibiting a wide range in size at maturity.

Journal of the Fisheries Research Board of Canada 30:607-616. 393

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Chen, Y. and H.H. Harvey. Growth, abundance, and food supply of White Sucker.

Transactions of the American Fisheries Society 124:262-271.

5. Doherty, C.A., R.A. Curry, and K.R. Munkittrick. 2010. Spatial and temporal

movements of White Sucker: Implications for use as a sentinel species.

Transactions of the American Fisheries Society 139(6):1818-1827.

6. McCormick, J.H., B.R. Jones, and K.E.F. Hokanson. 1977. White Sucker

(Catostomus commersonii) embryo development, and early growth and survival at

different temperatures. Journal of the Fisheries Research Board of Canada

34(7):1019-1025.

7. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

8. Reighard, J. 1920. The breeding behavior of the suckers and minnows. I. The

suckers. Biological Bulletin 38:1-32.

9. Reutter, J.M., and C.E. Herdendorf. 1976. Thermal discharge from a nuclear

power plant: predicted effects on Lake Erie fish. Ohio Journal of Science

76(1):39-45.

10. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

11. Twomey, K.A., K.L. Williamson, and P.C. Nelson. 1984. Habitat suitability index

models and instream flow suitability curves: white sucker. US Fish and Wildlife

Service FWS/OBS-82/10.64.

394

Mountain Sucker, Pantosteus jordani (Cope, 1874)

Etymology and Synonyms: Pantosteus = Panto, meaning “all”, and osteo, meaning

“bone”, likely referring to the joining of the parietal bones; jordani = honoring

Evermann’s friend and ichthyological colleague, David Starr Jordan.

Description: Body elongate, slender, cylindrical. Dorsally light dusky olive green to grey with dark mottling; laterally light olive to green; ventrally cream to white; fins unpigmented with no distinct markings. Head short, broad, conical. Snout blunt, long.

Eye moderately large. Mouth inferior, large; deep, lateral notches present on each corner.

Lips fleshy, papillose; lower lip lobes incompletely divided. Both jaws with inner cartilaginous ridge. Gill rakers 23-37 on external row; 31-51 on internal row of first gill arch.1 Dorsal fin with 8-13 rays, usually 10. Adipose fin absent. Caudal peduncle short.

Caudal fin slightly forked. Anal fin with 7 rays. Pelvic fin with 9 rays; insertion posterior to anterior end of dorsal fin. Pectoral fins rounded with ~15 rays. Lateral line complete with 75-92 small cycloid scales, crowded anteriorly. Intestine long, 4-6 times body length.3 Peritoneum black. Spawning males with well-defined burnt orange lateral stripe above a dark green to dark grey thin stripe extending from tip of snout to caudal fin base.

Spawning females with duller lateral stripe extending from operculum to anal fin.3 Males with tubercles on entire body including the enlarged anal fin rays. Females with tubercles only the dorsal and lateral sides of the head and body.3

Similar Species: Resembles the White and Longnose Sucker. White Sucker with a short snout slightly extending beyond the upper lip, 55-75 lateral line scales, no cartilaginous ridge on inner jaws, and no notches present on each corner of the mouth. Longnose 395

Sucker with more than 90 very small cycloid scales in lateral line, lower lip lobes completely divided, and no cartilaginous ridge on inner jaws.

Distribution and Habitat: Native to the western United States and Canada from southern Saskatchewan and Alberta in the north, south and west to eastern California, and east through the Intermountain range to western Nebraska and the Black Hills of South

Dakota.7,9 Historically occurred throughout the entire Black Hills region of South Dakota, however they are now constrained to the northern and southern portions of the Black

Hills, and are listed as a species of greatest conservation need in South Dakota.7,10 Not present in North Dakota. Although known to occur in large lakes, reservoirs, and rivers throughout its range, Mountain Sucker only inhabit cool water, low gradient streams with sand, gravel, or cobble substrates and moderate flow within the Black Hills.1,7 Density of the species increases when periphyton is abundant.5,8 Often associated with submerged vegetation and canopy cover. Optimal temperature 11-19 °C (51.8-66.2 °F).3 Critical thermal maximum for the Black Hills is 31.5 °C (88.7 °F), making the species much more tolerant of higher water temperatures than the co-occurring species of salmonids in the area.6 Fingerlings often associated with side channel habitats or deeper pools with little discharge.3 Able to tolerate moderate turbidity.

Reproduction: Spawning in the Black Hills is thought to be protracted. Males with breeding colors have been observed from early June through late August.2 Optimal water temperatures for spawning activity are 11-19 °C (51.8-66.2 °F).3,4 Spawning grounds consists of riffle areas below pools over gravel substrates in water roughly 18 cm (7.1 in) deep, with velocities of 12-15 cm/s.11 Eggs light yellow in color, average 1.77 mm (0.07 in) in diameter, and adhere to the substrate.11 Fecundity strongly correlated with size of 396 female.11 A group of 20 females averaging 162 mm (6.38 in) TL, had a mean fecundity of

2,087 eggs.11 Males known to reach sexual maturity at age 2-4; females at age 3-5.3 In the

Black Hills, the smallest mature male was 95 mm (3.7 in) TL, and the smallest mature female was 101 mm (4.0 in) TL.2

Age and Growth: Females generally are larger and live longer than males.3 In the Black

Hills, Mountain Sucker averaged 100 mm (3.93 in) TL during their fourth growing season.2 Average maximum TL in the Black Hills was 220 mm (8.66 in).2 Length at age from Whitewood and Elk Creeks in the Black Hills are recorded as: age-3, 100 mm (3.93 in) TL; age-4, 111 mm (4.37 in) TL, age-5, 133 mm (5.24 in) TL; age-6, 151 mm (5.94 in) TL.2 Growth of individuals in the Black Hills slower than growth rates in other parts of its range.2 Capable of reaching 222.5 mm (8.8 in) TL.3 Longevity 6-9 years.

Food and Feeding: Generalist benthic grazer. Predominantly herbivorous, mainly consuming periphyton and diatoms as well as smaller portions of aquatic insects and larvae such as dipterans. Being benthic feeders, sand and grit may also be found in the diet.4 Facilitate active feeding by scraping algae and benthic material off of hard substrates with cartilaginous ridges on the inside of both jaws.

Literature Cited:

1. Baxter, G.T., and M.D. Stone. 1995. Fishes of Wyoming. Wyoming Game and

Fish Department. Cheyenne, Wyoming.

2. Breeggemann, J.J., C.A. Hayer, J. Krause, L.D. Schultz, K.N. Bertrand, and

B.D.S. Graeb. 2014. Estimating the ages of Mountain Sucker Catostomus

platyrhynchus from the Black Hills: precision, maturation, and growth. Western

North American Naturalist 74(3):299-310. 397

3. Hauser, W.J. 1969. Life History of the Mountain Sucker, Catostomus

platyrhynchus, in Montana. Transactions of the American Fisheries Society,

98(2):209-215.

4. Moyle, P.B., R.M. Yoshiyama, J.E. Williams, and E.D. Wikramanayake. 1995.

Fish species of special concern in California, second edition. Department of

Wildlife and Fisheries Biology. University of California, Davis, Davis, California.

5. Schultz, L.D. 2011. Environmental factors associated with long-term trends of

Mountain Sucker populations in the Black Hills, and the assessment of their

thermal tolerance. Master’s thesis, South Dakota State University, Brookings,

South Dakota.

6. Schultz, L.D., and K.N. Bertrand. 2011. An assessment of the lethal thermal

maxima for Mountain Sucker. Western North American Naturalist 71(3):404-411.

7. Schultz, L.D., and K.N. Bertrand. 2012. Long term trends and outlook for

Mountain sucker in the Black Hills of South Dakota. The American Midland

Naturalist, 167(1):96-110.

8. Schultz, L.D., K.N. Bertrand, and B.D.S. Graeb. 2016. Factors from multiple

scales influence the distribution and abundance of an imperiled fish-mountain

sucker in the Black Hills of South Dakota, USA. Environmental Biology of Fishes

99:3-14.

9. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Resource Board of Canada, Bulletin 184. 966p. 398

10. SDGFP (South Dakota Department of Game, Fish, and Parks). 2006. South

Dakota Comprehensive Wildlife Conservation Plan. SDGFP, Wildlife Division

Report 2006-2008, Pierre.

11. Wydoski, R.G., and R.S. Wydoski. 2002. Age, growth, and reproduction of

Mountain Suckers in Lost Creek Reservoir, Utah. Transactions of the American

Fisheries Society 131(2):320-328.

399

Blue Sucker, Cycleptus elongates (Lesueur, 1817)

Etymology and Synonyms: Cycleptus = combination of the Greek words “kyklios” meaning round, and “leptes” meaning thin; elongates = elongate, referring to the body shape.

Description: Body elongate, slightly laterally compressed, terete shape. Dorsal side dark bluish gray, fading to light bluish gray on lateral sides; ventral side white; fins dark blue to dark gray. Head small. Snout elongate and pointed with rounded tip; extends beyond mouth. Mouth small, inferior. Teeth absent. Lips thick and heavily papillose. Eye small, placed laterally on anterior portion of head. Dorsal fin sickle-shaped with 28-36 rays; anterior rays elongated; spines absent. Caudal peduncle elongated with 19-20 rows of scales. Caudal fin moderately forked. Anal fin with 7-8 rays. Pelvic fins abdominal.

Pectoral fins falcate. Lateral line complete with 55-58 moderately sized cycloid scales in series. Spawning males darker in color with tubercles present on head, snout, body, and fins; spawning females light blue to tan with slight tubercles. Juveniles more elongate than adults, often with black spot present on lower caudal lobe.

Similar Species: May resemble carpsuckers (Carpiodes species) and buffalofishes

(Ictiobus species) due to the sickle-shaped dorsal fin, however the body of the Blue

Sucker is much more elongated and contains > 50 lateral line scales. The White Sucker has a similar mouth, however displays a short dorsal fin unlike the Blue Sucker.

Distribution and Habitat: Endemic to North America. Found within the Mississippi,

Missouri, and Rio Grande River basins and their main tributaries. Ranges from Montana to Mexico, and east to Pennsylvania.5 In the Dakotas, it has been reported in the Missouri

River drainage. Inhabits the lower main stems of the Big Sioux, James, and Vermillion 400

Rivers in South Dakota.2,5 Adults occur in deep waters with swift current in large rivers over firm substrate.1,6 Often associated with man-made structures such as wing dykes, or other flow controlling designs.6 Blue Sucker have reportedly declined throughout its range due to these flow controlling structures and river channelization, which alter spawning migrations and hydrology.1,7 Larval Blue Sucker occur in river backwaters along shallow (< 0.5 m deep), non-flowing shorelines in tributaries near the main channel.1,4 Low flow areas, especially those with islands, become important nursery areas for juvenile Blue Sucker.1 Larval and juvenile Blue Sucker are often found within tributaries with low flow, implying that tributaries are critical to their reproductive ecology.5,10

Reproduction: Little information is known about the reproductive behaviors of the Blue

Sucker. Spawning is known to take place within a short time period (10-28 days) from

April through early June when water temperatures reach roughly 14-18°C (57.2-

64.4°F).1,6,9 Males mature at roughly 4 years of age; females age-6.5,8 Spawning migration from large rivers to tributaries may occur, especially in years with high discharge.5,6,8

Tributaries with higher discharges may attract more spawning Blue Suckers than tributaries with lower flows.6 Spawning has been known to occur in swift, rocky riffles over bedrock or cobble substrate.6,9 Eggs opaque with yellow tint, adhesive, and average

2.2 mm (0.086 in) in diameter.6

Age and Growth: In the Mississippi River, individuals 16.0-39.0 mm (0.63-1.54 in) in length were estimated to be 12-42 days old; 13-20 mm (0.51-0.79 in) individuals were estimated to be 12-24 days old.1 Age-1 Blue Sucker in the Missouri River averaged 266 mm (10.47 in) TL; age-2 individuals averaged 323 mm (12.72 in).6 Growth is known to 401 be rapid during the juvenile stages (< age-5), and then taper off in the later years.5 In the

James River, Blue Sucker 374-717 mm (14.72-28.23 in) weighed 336-3026 g (0.74-6.67 lbs.); in the Big Sioux River, individuals 446-701 mm (17.56-27.60 in) weighed 508-

2730 g (1.12-6.02 lbs.).5 In the Kansas River, 500 mm was reached by age-7.3 Adults average 400-700 mm (15.75-27.5 in).5,3 Males often smaller than females of the same age.6 Can live to 16 years although most have a lifespan of 7-11 years.3

Food and Feeding: Adults feeds primarily on immature aquatic insects such as mayflies, midges, black flies, and caddisflies. Also consumes zooplankton, nematodes, fingernail clams, filamentous algae, and leaf litter.1,6 Larval Blue Suckers have a diet mainly composed of chironomids, copepods, and cladocerans.1 Both aquatic and terrestrial invertebrates within backwaters around islands are consumed by juvenile Blue Sucker and are most likely very critical to the early survival of the species.1

Literature Cited:

1. Adams, S.R., M.B. Flinn, B.M. Burr, M.R. Whiles, and J.E. Garvey. 2006.

Ecology of larval blue sucker (Cycleptus elongates) in the Mississippi River.

Ecology of Freshwater Fish 15:291-300.

2. Dieterman D. and C.R. Berry, Jr. 1995. The distribution and relative abundance of

fishes in the Big Sioux River, South Dakota. Progress Report 95-9, Department of

Game, Fish, and Parks, Pierre.

3. Eitzmann, J.L., A.S. Makinster, and C.P. Paukert. 2007. Distribution and growth

of blue sucker in a Great Plains river, USA. Fisheries Management and Ecology

14:255-262. 402

4. Fisher, S.J. and D.W. Willis. 2000. Observations of age-0 blue sucker, Cycleptus

elongates, utilizing an upper Missouri River backwater. Journal of Freshwater

Ecology 15:425-427.

5. Morey, N.M. and C.R. Berry Jr. 2003. Biological Characteristics of the Blue

Sucker in the James River and the Big Sioux River, South Dakota. Journal of

Freshwater Ecology 18(1): 33-41.

6. Moss, R.E., J.W. Scanlan, and C.S. Anderson. 1983. Observations on the natural

history of the blue sucker (Cycleptus elongatus Le Sueur) in the Neosho River.

American Midland Naturalist 109 (1): 15-22.

7. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

8. Rupprecht, R.J. and L.A. Jahn. 1980. Biological notes on blue suckers in the

Mississippi River. Transactions of the American Fisheries Society 109:323-326.

9. Vokoun, J.C., T.L. Guerant, and C.F. Rabeni. 2003. Demographics and

chronology of a spawning aggregation of blue sucker (Cycleptus elongates) in the

Grand River, Missouri, USA. Journal of Freshwater Ecology 18:567-575.

10. Young, B.A., T.L. Welker, M.L. Wildhaber, C.R. Berry, and D. Scarnecchia,

editors. 1998. Population structure and habitat use of benthic fishers along the

Missouri and Lower Yellowstone Rivers. 1997 Annual Report of Missouri River

Benthic Fish Study PD-95-5832 to the U.S. Army Corps of Engineers and the

U.S. Bureau of Reclamation.

403

Smallmouth Buffalo, Ictiobus bubalus (Rafinesque, 1818)

Etymology and Synonyms: Ictiobus = ichthys, meaning “fish”, and bous, meaning “ox”, likely referring to the species common name or robust appearance; bubalus = “wild ox”, or “buffalo”.

Description: Body robust, deep, slightly laterally compressed with a steep, “humped” profile from head to anterior end of dorsal fin. Dorsally bronze, olive green; laterally light olive; ventrally light yellow-gray to white; fins gray to light brown without markings.

Head small, conical. Subopercle with rounded outline on lower corner. Snout blunt. Eye small, positioned anteriorly on head. Mouth small, subterminal. Barbels absent. Lips thick, fleshy with deep grooves present. Teeth absent on jaws; roughly 130-195 small, comblike pharyngeal teeth per gill arch. Gill rakers 30-35 on first arch. Dorsal fin long, falcate with 25-31 rays. Adipose fin absent. Caudal peduncle short, thick. Caudal fin moderately forked with slightly rounded lobes. Anal fin with 8-10 rays. Pelvic fin abdominal with 9-11 rays. Pectoral fin with straight distal end. Lateral line complete with

35-39 cycloid scales in series. Spawning males develop fine tubercles on head.

Similar Species: Closely resembles the Black and Bigmouth Buffalo. Black Buffalo with a slightly compressed, more slender body, large head, also with a small, subterminal to inferior mouth with thick, deeply grooved lips. Bigmouth Buffalo with a less deep body, a large head leading to a rounded profile to the anterior end of the dorsal fin, less steep than the Smallmouth Buffalo, and thin lips with only the lower lip having shallow grooves present. Common Carp with 2 fleshy barbels present on each side of mouth.

Carpiodes species, such as carpsucker and Quillback, have an angled outline on the lower corner of the subopercle, unlike Ictiobus species. 404

Distribution and Habitat: Native throughout the Mississippi-Missouri River basin from

Montana in the west, east to Pennsylvania, and south to the Gulf of Mexico. Occurs in the

Missouri, James, Big Sioux, Vermillion and Red River of the North in the Dakotas. The twelve records from the Red River of the North may be misidentification but are unable to be verified as specimens were not vouchered from those collections. Adults occur most often in medium to large sized rivers in areas with low velocity such as backwaters, oxbows and pools. Also inhabits reservoirs and lakes. Less frequent in small streams. Prefers clean, clear, warm water with firm substrates and abundant aquatic vegetation. Migrates to shallow waters in the spring and summer, and retreats to deeper water in fall and winter.3,5 Known to inhabit deeper depths than the Bigmouth Buffalo. Capable of withstanding areas of moderate current for short periods of time.3 Fry and juveniles frequent warm, shallow, vegetated backwaters or embayments with water velocity <20 cm/sec.3 Can tolerate turbid waters (>100 NTU), but to a lesser extent than Bigmouth

Buffalo. Optimum pH is 6.5-8.5.3

Reproduction: Spawning initiated and most successful with rising water levels.5,10

Spawning takes place from March to April, but may continue until September.5 Optimal spawning temperature is 19.1-27.5 °C (6.6.4-81.5 °F), with peak spawning occurring at

22-26.5 °C (71.6-79.7 °F).5 Age at sexual maturity variable, and may be a function of size rather than age.5 In Lewis and Clark Lake, males reach sexual maturity at age 7-9; females age 10-11.11 Early in the season, males move upstream and position themselves near shallow shoals, backwaters, or within pools roughly 1.2-3.1 m (4-10 ft.) deep, over or near vegetation where they congregate with ripe females.3,5 No nest is created, and no parental care is given. Fecundity dependent on size; an age-15 female contained 525,500 eggs.5 Eggs broadcast, demersal, adhesive to bottom substrate or vegetation, and roughly 405

1.5 mm (0.05 in) in diameter.5 Hatching occurs within 4-14 days depending on water temperature.

Age and Growth: Newly hatched larvae roughly 6 mm (0.24 in) TL, and 105 mm (4.13 in) TL by the first summer.13 Growth rates better in clear to slightly turbid water.3,8,12

Seasonal growth initiated when water temperatures reach 18 °C (64.4 °F).9 Mean lengths- at-age from Wisconsin are reported as: age-1, 112 mm (4.41 in) TL; age-2, 245 mm (9.65 in) TL; age-3, 346 mm (13.62 in) TL; age-4, 391 mm (15.39 in) TL; age-5, 428 mm

(16.85 in) TL; age-6, 462 mm (18.19 in) TL.1 Capable of reaching >762 mm (30 in) TL, and 14-18 kg (30-40 lbs.). Longevity 15-20 years.5

Food and Feeding: Opportunistic planktivore. In Lewis and Clark Lake, fry primarily consume microcrustacea such as Copepods and Cladocerans, in shallow areas with low flow.7 Young adult and adult diets mainly consist of attached algae and zooplankton, especially Copepods and Cladocerans such as Cyclops, Bosmina, and Daphnia.7 A large palatal organ situated in the pharynx assists the species with selective food retention.2,6

Chlorophyta, insect larvae, detritus, and sand found in diets indicate that the species likely feeds along the bottom near the shoreline.7 Mollusks, insect larvae and insect eggs have also been reported in diets.4

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Doosey, M.H., and H.L. Bart. 2011. Morphological variation of the palatal organ

and chewing pad of Catostomidae (Teleostei: Cypriniformes). Journal of

Morphology 272:1092-1108. 406

3. Edwards, E.A., and K. Twomey. 1982. Habitat suitability index models:

Smallmouth Buffalo. U.S. Department of the interior, Fish and Wildlife Service.

FWS/OBS-82/10.13.

4. Forbes, S.A., and R.E. Richardson. 1920. The fishes of Illinois. Illinois State

Journal Company, State Printers, Springfield.

5. Jester, D.B. 1973. Life history, ecology, and management of the Smallmouth

Buffalo, Ictiobus bubalus (Rafinesque), with reference to Elephant Butte Lake,

New Mexico. New Mexico State University Agricultural Experiment Station

Research Report 261.

6. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

7. McComish, T.S. 1967. Food habits of Bigmouth and Smallmouth Buffalo in

Lewis and Clark Lake and the Missouri River. Transactions of the American

Fisheries Society 96(1):70-74.

8. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

9. Shields, J.T. 1957. Report of fisheries investigations during the fourth year of

impoundment of Fort Randall Reservoir, South Dakota, 1956. South Dakota

Department of Game, Fish, and Parks Dingell-Johnson Project. F-1-R-6:1-60.

10. Walburg, C.H. 1976. Changes in the fish populations of Lewis and Clark Lake,

1956-74, and their relation to water management and the environment. U.S. Fish

and Wildlife Service, Res. Report 79. 34pp. 407

11. Walburg, C.H., and W.R. Nelson. 1966. Carp, River Carpsucker, Smallmouth

Buffalo, and Bigmouth Buffalo in Lewis and Clark Lake, Missouri River. U.S.

Department of Interior, Fish and Wildlife Service Res. Report 69. 30pp.

12. Willis, D.W. 1978. Investigations of population structure and relative abundance

of year-classes of buffalo fishes, Ictiobus spp., in Lake Sakakawea, North Dakota.

M.S. Thesis, University of North Dakota, Grand Forks.

13. Wrenn, W.B., and B.G. Grinstead. 1971. Larval development of the Smallmouth

Buffalo, Ictiobus bubalus. Journal of the Tennessee Academy of Science 46:117-

120.

408

Bigmouth Buffalo, Ictiobus cyprinellus (Valenciennes, 1844)

Etymology and Synonyms: Ictiobus = ichthys, meaning “fish”, and bous, meaning “ox”, likely referring to the species common name or robust appearance; cyprinellus = Greek meaning for “carp”.

Description: Body robust, moderately deep, slightly laterally compressed, with distinct rounded profile from head to anterior end of dorsal fin. Dorsally dark gray to olive- brown; laterally dusky brown to olive; ventrally light gray to white; fins dark gray to black without markings; black and orange markings on dorsal and head typically present and more intensified in older individuals. Head large. Subopercle with rounded outline on lower corner. Snout blunt with slight depression between eyes. Eye small, positioned anteriorly on head. Mouth large, terminal and oblique. Barbels absent. Lips thin, fleshy, smooth; lower lip with shallow grooves present. Teeth absent on jaws; roughly 130-170 short, comblike pharyngeal teeth per gill arch. Gill rakers long, 40-45 on first arch.

Dorsal fin long, falcate with 24-32 rays. Adipose fin absent. Caudal peduncle short, thick.

Caudal fin moderately forked with slightly rounded lobes. Anal fin with 8-10 rays. Pelvic fin abdominal with 10-11 rays. Pectoral fin with straight distal end. Lateral line complete with 34-40 cycloid scales in series. Spawning males develop small tubercles on head, nape of neck and fin rays.

Similar Species: Closely resembles the Smallmouth and Black Buffalo. Smallmouth

Buffalo with a deeper body, and a small, conical head creating a steep, or “humped” profile from the head to anterior end of dorsal fin. Smallmouth Buffalo also have a small, subterminal mouth and thick lips with deep grooves present. Black Buffalo with a slightly compressed, more slender body, also with a small, subterminal to inferior mouth with 409 thick, deeply grooved lips. Common Carp with 2 fleshy barbels present on each side of mouth. Carpiodes species, such as River Carpsucker and Quillback, have an angled outline on the lower corner of the subopercle, unlike Ictiobus species.

Distribution and Habitat: Native throughout the Mississippi-Missouri River basin, as well as the lower Great Lakes and Hudson Bay drainages. Occurs in the Missouri, James,

Big Sioux, Vermillion and Red Rivers in the Dakotas. Inhabits low gradient, medium to large sized rivers in areas with low flow. Also well adapted to reservoirs and lakes.8

Often associated with silt, sand, or gravel substrates, and areas with abundant vegetation.5

Migrates upstream in the spring to spend summers in shallow, warm waters, and migrates back to deeper water in the fall to overwinter.2,3,5 Prefers shallow water, 1.2-3.1 m (4-10 ft.), and is sometimes seen just below the surface in schools on warm summer days.8,13

Optimum water temperature 31-34°C (87.8-93.2°F).5,7 Fry and juveniles frequent shallow bay areas with little to no flow.8 More tolerant of turbidity than Smallmouth Buffalo or

Black Buffalo.6 Able to withstand low oxygen levels and high water temperatures.1

Optimum pH 6.5-8.5.5

Reproduction: Spawning takes place from April to June, or when water temperatures reach 14-18°C (57.2-64.4°F), in backwaters or shallow areas near aquatic vegetation such as cattails.3,5,8 Spawning tends to cease when water temperature reaches 26.7°C

(80.06°F).5 Annual spawner, unless suitable habitat is not available, in which case spawning may be skipped.12 In Lewis and Clark Lake, males reach sexual maturity at age

7-8; females age 8-9.13 Spawning ritual consists of a single female accompanied by two or more males splashing near the surface until the group sinks to the bottom where the eggs are fertilized and released.4 After activity ceases, the fish migrate up to 0.8-1.6 km 410

(0.5-1.0 mi) back to the mainstream channel or original location.1 No nest is created, and no parental care is given. Fecundity dependent on size, but can reach 750,000 eggs per female.8 Eggs adhesive, roughly 1.2-1.8 mm (0.05-0.07 in) in diameter.8 Hatching occurs within 8-14 days depending on water temperature.

Age and Growth: Largest of the Ictiobus species. Growth variable with location and rapid within first few years of life with individuals reaching 50-150 mm (1.97-5.12 in)

TL during their first summer. Mean lengths-at-age from Wisconsin are reported as: age-

1, 131 mm (5.16 in) TL; age-2, 255 mm (10.04 in) TL; age-3, 335 mm (13.19 in) TL; age-4, 393 mm (15.47 in) TL; age-5, 453 mm (17.83 in) TL; age-6, 508 mm (20 in) TL; age-7, 561 mm (22.09 in) TL; age-8, 614 mm (24.17 in) TL.1 Capable of reaching 1,250 mm (49.21 in) TL, and >36 kg (79.4 lbs.).5,9 Longevity 112 years, making them to date, the oldest age-validated freshwater teleost fish.9

Food and Feeding: Opportunistic planktivore. Long gill rakers assist the species in filtering small, planktonic prey throughout the water just above the bottom substrate. Fry known to feed in shallow bays and consume rotifers, small crustaceans such as copepods and cladocerans (especially Daphnia), and benthic organisms such as tendipedid pupae and larvae.10,11 Young adult and adult diets primarily consist of copepods and cladocerans

(especially Daphnia), and midge larvae.10,11 Cyanobacteria such as Anacytis are known to be important food sources from late summer to fall.11 Occasionally known to consume aquatic beetles and small mollusks.8

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press,

Madison. 411

2. Beckman, L.G., and J.H. Elrod. 1971. Apparent abundance and distribution of

young-of-the-year fishes in Lake Oahe, 1965-69. Pages 333-347 in G.E. Hall

(ed.). Reservoir fisheries and limnology. American Fisheries Society Special

Publication 8.

3. Benson, N.G. 1980. Effects of post impoundment shore modification on fish

populations in Missouri River reservoirs. U.S. Department of Interior, Fish and

Wildlife Service Research Report 80. 32pp.

4. Burr, B.M., and R.C. Heidinger. 1983. Reproductive behavior of the Bigmouth

Buffalo Ictiobus cyprinellus in Crab Orchard Lake, Illinois. American Midland

Naturalist 110:220-221.

5. Edwards, E.A. 1983. Habitat suitability index models: Bigmouth Buffalo. U.S.

Department of the Interior, Fish and Wildlife Service. FWS/OBS-82/10.34.

6. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

7. Gammon, J.R. 1973. The effect of thermal input on the populations of fish and

macroinvertebrates in the Wabash River. Purdue University Water Resource

Center, Lafayette, Indiana. Technical Report 32. 106pp.

8. Johnson, R.P. 1963. Studies on the life history and ecology of the Bigmouth

Buffalo, Ictiobus cyprinellus (Valenciennes). Journal of Fisheries Research Board

of Canada 20(6):1397-1429.

9. Lackman, A.R., A.H. Andrews, M.G. Butler, E.S. Bielak-Lackman, and M.E.

Clark. 2019. Bigmouth Buffalo, Ictiobus cyprinellus sets freshwater teleost record 412

as improved age analysis reveals centenarian longevity. Communications Biology

197 (2):1-14.

10. McComish, T.S. 1967. Food habits of Bigmouth and Smallmouth Buffalo in

Lewis and Clark Lake and the Missouri River. Transactions of the American

Fisheries Society 96(1):70-74.

11. Starostka, V.J., and R.L. Applegate. 1970. Food selectivity of Bigmouth Buffalo,

Ictiobus cyprinellus, in Lake Poinsett, South Dakota. Transactions of the

American Fisheries Society 99(3):571-576.

12. Walburg, C.H. 1976. Changes in the fish populations of Lewis and Clark Lake,

1956-74, and their relation to water management and the environment. U.S. Fish

and Wildlife Service Research Report 79. 34pp.

13. Walburg, C.H., and W.R. Nelson. 1966. Carp, River Carpsucker, Smallmouth

Buffalo, and Bigmouth Buffalo in Lewis and Clark Lake, Missouri River. U.S.

Department of Interior, Fish and Wildlife Service Research Report 69. 30pp.

413

Black Buffalo, Ictiobus niger (Rafinesque, 1819)

Etymology and Synonyms: Ictiobus = ichthys, meaning “fish”, and bous, meaning “ox”, likely referring to the species common name or robust appearance; niger = black.

Description: Body slightly robust, elongate, and slightly compressed with slightly rounded profile from head to anterior end of dorsal fin. Dorsally black to dark olive-gray; laterally dusky brown to olive; ventrally light gray to white; fins dark gray to black without markings. Head large. Subopercle with rounded outline on lower corner. Snout blunt. Eye small, positioned anteriorly on head. Mouth small, subterminal to inferior.

Lips thick, with deep grooves present. Teeth absent on jaws; roughly 195 short pharyngeal teeth per gill arch. Gill rakers 30-35 on first arch. Dorsal fin long, falcate with

25-31 rays. Adipose fin absent. Caudal peduncle short, thick. Caudal fin moderately forked with slightly rounded lobes. Anal fin with 8-9 rays. Pelvic fin abdominal with 9-

11 rays. Pectoral fin with slightly rounded distal end. Lateral line complete with 36-39 cycloid scales in series. Spawning males develop small tubercles on head, anterior scales, and pelvic fins.

Similar Species: Closely resembles the Smallmouth and Bigmouth Buffalo. Smallmouth with a deeper, slightly laterally compressed body, and a small conical head creating a steep, or “humped” profile from the head to anterior end of dorsal fin. Smallmouth

Buffalo also have a small, subterminal mouth and thick lips with deep grooves present.

Bigmouth Buffalo with a slightly more robust body, more exaggerated profile or “hump” from head to anterior end of dorsal fin, and a large, terminal mouth with thin lips.

Common Carp with 2 fleshy barbels present on each side of mouth. Carpiodes species, 414 such as River Carpsucker and Quillback, have an angled outline on the lower corner of the subopercle, unlike Ictiobus species.

Distribution and Habitat: Least common Ictiobus species in its range. Native throughout the lower Great Lakes and Mississippi-Missouri River basins from southeastern South Dakota, east to Michigan, and south to Louisiana. Occurs in the

Missouri, James, and Big Sioux rivers in the Dakotas, but to a much lesser extent than

Bigmouth or Smallmouth Buffalo. Inhabits medium to large sized rivers, streams, and reservoirs. Often found in sloughs, and backwaters near deep, swift riffles. Known to prefer greater water velocities than the Bigmouth and Smallmouth Buffalo.1 Also found in shallow waters with bedrock or gravel substrate.2 Less tolerable of turbidity than

Bigmouth Buffalo.

Reproduction: Little is known about the reproductive biology of the Black Buffalo compared to other Ictiobus species, but behavior thought to be similar to Smallmouth

Buffalo.4 Annual spawner. Sexual maturity reached at age-2 in southern populations.1,8

Spawning is presumed to take place April-May. In riverine habitat, adults known to migrate from deeper water within the main channel to shallow backwaters, runs, and pools.4,9 Spawning ritual consists of a single female accompanied by two or more males.9

Males swim alongside the female and bump her near the surface to induce the release of her eggs.9 This behavior is known to create splashing.9 Eggs demersal and adhesive.

Age and Growth: Adults average roughly 381-762 mm (15-30 in) TL, and 4.5-6.8 kg

(10-15 lbs). Mean lengths-at-age from Kansas are reported as: age-1, 310 mm (12.2 in)

TL; age-2, 368 mm (14.5 in) TL; age-3, 447 mm (17.6 in) TL; age-4, 543 mm (21.4 in) 415

TL; age-5, 627 mm (24.7 in) TL.5,6 Capable of reaching 800 mm (31.5 in) TL, and 9.07-

13.61 kg (20-30 lbs). Longevity >20 years.

Food and Feeding: Little information exists on food and feeding habits of Black

Buffalo, but it is known to be similar to Bigmouth Buffalo. Opportunistic feeder. Highly dependent on benthic organisms.7 Primarily feeds along the bottom, stirring up sediments in search for prey items. Feeding on plankton throughout the water column seems unlikely due to the positioning of the mouth.4 Adults mainly consume diatoms, cyanobacteria, crustaceans, insects and detritus.4 In Arizona, juvenile Asian clams made up the majority of the adult diets.4 A large palatal organ situated in the pharynx assists the species with selective food retention.3,6

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Cross, F.B.1967. Handbook of fishes in Kansas. University of Kansas Museum of

Natural History, Miscellaneous Publication Number 45:1-357.

3. Doosey, M.H., and H.L. Bart. 2011. Morphological variation of the palatal organ

and chewing pad of Catostomidae (Teleostei: Cypriniformes). Journal of

Morphology 272:1092-1108.

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

5. Greer, J.K., and F.B. Cross. 1956. Fishes of El Dorado City Lake, Butler County,

Kansas. Transactions of the Kansas Academy of Science 59:358-363.

6. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence. 416

7. Minckley, W.L., J.E. Johnson, J.N. RInne, and S.E. Willoughby. 1970. Foods of

buffalofishes, genus Ictiobus, in central Arizona reservoirs. Transactions of the

American Fisheries Society 99:333-342.

8. Perry, W.G. 1976. Black and bigmouth buffalo spawn in brackish water ponds.

The Progressive Fish-Culturist 38(2):81.

9. Piller, K.R., H.L. Bart Jr., and J.A. Tipton. 2003. Spawning in the black buffalo,

Ictiobus niger (Cypriniformes: Catostomidae). Ichthyological Explorations of

Freshwater 14:145-150.

417

Silver Redhorse, Moxostoma anisurum

Etymology and Synonyms: Moxo = Greek for myxo, meaning, “to suck”, stoma = Greek for “mouth”; anisurum = “unequal tail”, referring to the asymmetry of the upper and lower lobes.

Description: Body fusiform to slightly dorso-ventrally flattened, elongate. Dorsally dark gray to olive; laterally silver, scales generally lacking dark spots at anterior base; ventrally white; dorsal and caudal fins dark gray in color; anal, pelvic and pectoral fins clear to light red-orange. Head moderately large. Snout blunt. Eye moderately large, placed laterally on upper portion of head. Mouth large, inferior; jaws do not extend backward to anterior end of eye. Lips plicate; halves of lower lip meet at an acute angle; majority of lower lips with plicae broken into individual papillae; lower lip thicker than upper lip. Teeth absent on jaws. Pharyngeal teeth long, slender. Gill rakers 20-32. Dorsal fin with 14-17 rays, straight to slightly convex distal end. Adipose fin absent. Caudal peduncle short, thick. Circumpeduncular scales in row usually 12. Caudal fin forked, upper lobe slightly more elongate and pointed. Anal fin with 7 rays. Pelvic fins with 9 rays; insertion posterior to insertion of dorsal fin. Pectoral fins with 16-19 rays. Lateral line complete with 41-46 large cycloid scales. Spawning males develop tubercles on head, body, lower lobe of caudal fin, and anal fin. Juveniles similar in appearance to adults.

Similar Species: Closely resembles other redhorse (Moxostoma) species. Shorthead,

River, and Greater Redhorse all display a red caudal fin and have dark spots present on anterior base of scales on lateral sides. Further, Shorthead Redhorse posterior margin of lower lip forms a straight line. River Redhorse lower lips plicae, but not broken into 418 individual papillae. Greater Redhorse with 13-15 dorsal fin rays, posterior margin of lower lip forming an obtuse angle, and typically 16 scales in circumpeduncular row.

Golden Redhorse posterior margin of lower lip forms a nearly straight line to obtuse angle, and have 39-42 lateral line scales in series.

Distribution and Habitat: Native to Canada from Alberta to Quebec, and in the United

States from the Red River of the North in eastern North Dakota, west throughout the upper and mid central Mississippi River, Great Lakes-St. Lawrence River, and Ohio

River drainages to Vermont in the east, and south to Alabama and the lower Missouri

River.5,6,8 Found within the Bois de Sioux River of the Red River drainage in South

Dakota. Inhabit small to large rivers, lakes and impoundments.10 In riverine habitats the species is often found within long, deep pools or backwater habitats with slow current over silty to firm substrate.4,7,10 Avoids high gradient streams with increased levels of turbidity.1 During their first year, young individuals inhabit smaller streams with slow water velocity, often near overhanging vegetation and stream banks to seek shelter from predators.1,3,7

Reproduction: Information regarding spawning ritual, egg description and hatching time is lacking. Records have indicated that Silver Redhorse may make small spawning migrations.11 Spawning in the northern portion of the species range likely takes place mid-April to early May.7,11 Individuals from the Des Moines River in Iowa reached peak spawning conditions when water temperatures reached 13.33°C (56°F) during the first week of May.7 In Alabama, spawning occurs late March to early April when water temperatures reach 14-15°C (57.2-59°F).2 Sexual maturity reached at age-5.7 Spawning takes place in the main channel of rivers and streams at depths around 0.3-0.9 m (0.98- 419

2.95 ft.) over coarse substrate.7 In Iowa, fecundity ranged 14,910 eggs from an age-5,

337.82 mm (13.3 in) TL female, to 36,340 eggs from an unknown age, 490.22 mm (19.3 in) TL female.7 Eggs 1.1-2.2 mm (0.04-0.09 in) in diameter.1

Age and Growth: Largest annual growth rates occur from late July to early September.1,4

Mean lengths-at-age from the Des Moines River in Iowa are reported as: age-1, 109.22 mm (4.30 in) TL; age-2, 194.56 mm (7.66 in) TL; age-3, 270.76 mm (10.66 in) TL; age-

4, 328.42 mm (12.93 in) TL; age-5, 379.22 mm (14.93 in) TL; age-6, 431.8 mm (17.00 in) TL; age-7, 470.15 mm (18.51 in) TL; age-8, 504.95 mm (19.88 in) TL; age-9, 513.08 mm (20.20 in) TL.7 Capable of reaching 675 mm (26.57 in) TL.9 Longevity 25 years.9

Food and Feeding: Information on diet and feeding habits of Silver Redhorse is lacking.

Adults consume immature aquatic insects such as Chironomids, and Ephemeroptera and

Trichoptera larvae, as well as filamentous algae and detritus.7,11

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

3. Harlan, J.R., and E.B. Speaker. 1956. Iowa fish and fishing. Iowa Conservation

Committee. 377pp.

4. Jenkins, R.E. 1970. Systematic studies of the catostomid fish Moxostomatini.

Ph.D. Dissertation. Cornell University, Ithaca, New York.

5. Jenkins, R.E. 1980. Moxostoma anisurum. Pages 409-410 in Atlas of North

American freshwater fishes (D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, 420

D.E. McAllister, and J.R. Stauffer, Jr., editors). North Carolina State Museum of

Natural History, Raleigh.

6. McAllister, C.T., W.C. Starnes, H.W. Robinson, R.E. Jenkins, and M.E. Raley.

2009. Distribution of the Silver Redhorse, Moxostoma anisurum (Cypriniformes:

Catostomidae), in Arkansas. The Southwestern Naturalist 54:514-518.

7. Meyer, W.H. 1962. Life history of three species of redhorse (Moxostoma) in the

Des Moines River, Iowa. Transactions of the American Fisheries Society 91:412-

419.

8. Page, L.M., and B.M. Burr. 1991. A field guide to freshwater fishes: North

America north of Mexico. Houghton Mifflin, Boston, Massachusetts.

9. Reid, S.M. 2007. Comparison of scales, pectoral fin rays, and opercles for age

estimation of Ontario Redhorse, Moxostoma, species. Canadian Field-Naturalist

121:29-34.

10. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

11. Smith, C.G. 1977. The biology of three species of Moxostoma (Pisces:

Catostomidae) in Clear Creek, hocking and Fairfield counties, Ohio, with

emphasis on the Golden Redhorse, M. erythrurum (Rafinesque). Ph.D.

Dissertation, Ohio State University, Columbus, Ohio.

421

River Redhorse, Moxostoma carinatum (Cope, 1870)

Etymology and Synonyms: Moxo = Greek for myxo, meaning, “to suck”, stoma = Greek for “mouth”; carinatum = “keeled”.

Description: Body fusiform, elongate, moderately robust. Dorsally olive to bronze; laterally golden yellow, olive to silver, scales with dark spots usually present at anterior base; ventrally white; dorsal fin dusky red; caudal fin red; anal, pelvic, and pectoral fins orange-red in color. Head moderately large. Snout blunt, almost square in shape. Eye moderately large, placed laterally on upper portion of head. Mouth large, inferior; jaws do not extend backward to anterior end of eye. Lips plicate; posterior margin of lower lips nearly straight or forming a slightly obtuse angle; lower lip plicae not dissected into individual papillae; lower lip thicker than upper lip. Teeth absent on jaws. Pharyngeal teeth large, molariform. Gill rakers 20-31. Dorsal fin with 13-15 rays, straight to slightly concave distal end. Adipose fin absent. Caudal peduncle slender, slightly elongate.

Circumpeduncular scales rows usually 12. Caudal fin forked, upper lobe slightly more elongate and pointed. Anal fin with 7 rays. Pelvic fins with 9 rays; insertion posterior to insertion of dorsal fin. Pectoral fins with 15-17 rays. Lateral line complete with 41-47 large cycloid scales in series. Spawning males develop large tubercles on snout, head, and opercle, as well as small tubercles on body, caudal and anal fins. Larger spawning females develop tubercles on the anal fin. Juveniles similar in appearance to adults.

Similar Species: Closely resembles other redhorse (Moxostoma) species. Silver and

Golden Redhorse both display a slate colored caudal fin, and generally lack dark spots on anterior base of scales on lateral sides. Further, Silver Redhorse posterior margin of lower lip forms an acute angle. Golden Redhorse posterior margin of lower lip forms a nearly 422 straight line or an obtuse angle, and have 39-42 lateral line scales. Shorthead Redhorse with a falcate to concave distal end of dorsal fin and posterior margin of lower lip forms a straight line. Greater Redhorse with a slightly convex distal end of dorsal fin, posterior margin of lower lip meeting at an obtuse angle, and usually 16 scales in circumpeduncular row.

Distribution and Habitat: Wide but discontinuous native distribution that extends throughout the eastern United States and southeastern Canada throughout the Great

Lakes, central Mississippi and Gulf slope drainages.11 Populations within the native historic range have experienced substantial declines over the past century, likely due to habitat loss.3,5,7 Inhabits deep riffle-runs areas >1.5 m (4.92 ft.) of medium to large sized rivers and lower portions of their main tributaries with clear water and moderate to rapid current >0.4 m/s, but may also occupy reservoirs.2,3,8 Occur over course substrates such as gravel, cobble or boulder.3 Movement patterns known to vary by season, with the largest ranges and highest displacements occurring during spring, and the smallest and lowest displacements taking place in summer.3 Areas with faster current are used more often during winter and spring.3 Intolerant to sedimentation and pollution, and negatively impacted by habitat fragmentation and flow regulation.3,8,12

Reproduction: In the northern part of the species ranges, spawning takes place in late

May through early June when water temperatures reach 17.5-19.0°C (63.5-66.2°F), with individuals reaching the post-spawning stage at 20°C (68°F).4 Sexual maturity reached around age-5.10 Males inhabit spawning grounds prior to females.4,6,9 Males in Alabama have been observed sweeping coarse substrate with their bodies and caudal fin to form redds, however this behavior has not been documented in other locations.4,6 Spawning 423 takes place day and night in the main channels of rivers and streams in areas <2 m (6.56 ft.) deep, within riffles over gravel substrate.3,4,11 Observed spawning events ranged 3.0-

13.5 seconds long, and occur over a short spawning season of 5-8 days.6,11 Estimated fecundity 9,000-22,000 eggs.4 Eggs yellow to yellow-orange in color, adhesive and average 2.8 mm (0.11 in) in diameter.4 Eggs incubated in a laboratory hatched within 3-4 days at a water temperature of 22.2°C (71.96°F).1,6

Age and Growth: One of the larger species of Moxostoma, often reaching >5 kg (11.02 lbs.).2,7 Information regarding mean age-at length is lacking. Majority of growth occurs within the first 5 years of life, with growth rates slowing at roughly age-8.4 Females tend to be larger than males.9 Capable of reaching 798 mm (31.42 in) TL.4 Longevity 28 years.4

Food and Feeding: Benthivore. Enlarged pharyngeal arches and molariform pharyngeal teeth assist the species when consuming mollusks such as mussels and snails, which comprise a large part of their diet.8 Aquatic insect larvae such as Emphemeropterans,

Chironomids, and Trichopterans are also known to be consumed.6

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Beckman, D.W., and C.A. Hutson. 2012. Validation of aging techniques and

growth of the River Redhorse, Moxostoma carinatum, in the James River,

Missouri. The Southwestern Naturalist 57:240-247.

3. Butler, S.E., and D.H. Wahl. 2017. Movements and habitat use of River Redhorse

(Moxostoma carinatum) in the Kankakee River, Illinois. Copeia 105:734-742. 424

4. Campbell, B.G. 2001. A study of the River Redhorse, Moxostoma carinatum

(Pisces: Catostomidae), in the tributaries of the Ottawa River near Canada’s

National Capital and in a tributary of Lake Ontario, the Grand River, near

Cayuga, Ontario. M.S. Thesis, University of Ottawa, Ottawa.

5. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

6. Hackney, P.A., W.M. Tatum, and S.L. Spencer. 1967. Life history of the River

Redhorse, Moxostoma carinatum (Cope), in the Cahaba River, Alabama, with

notes on the management of the species as a sport fish. Proceedings of the 21st

annual Conference of the Southeast Association of the Game Fisheries

Commission.

7. Jenkins, J.E., and N.M. Burkhead. 1993. Freshwater fishes of Virginia. American

Fisheries Society, Bethesda, Maryland.

8. Jenkins, R.E. 1970. Systematic studies of the catostomid fish tribe

Moxostomatini. Doctoral dissertation, University of Michigan, Ann Arbor.

9. Reid, S.M. 2006. Timing and demographic characteristics of redhorse spawning

runs in three Great Lakes basin rivers. Journal of Freshwater Ecology 21:249-258.

10. Ross, S.T. 2001. The inland fishes of Mississippi. Mississippi Department of

Wildlife, Fisheries and Parks, and University Press of Mississippi, Jackson.

11. Straight, C.A., C.R. Jackson, B.J. Freeman, and M.C. Freeman. 2015. Diel

patterns and temporal trends in spawning activities of Robust Redhorse and River

Redhorse in Georgia, assessed using passive acoustic monitoring. Transactions of

the American Fisheries Society 144:563-576. 425

12. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

426

Golden Redhorse, Moxostoma erythrurum (Rafinesque, 1818)

Etymology and Synonyms: Moxo = Greek for myxo, meaning, “to suck”, stoma = Greek for “mouth”; erythro = “red”, urum = “tail”, possibly referring to the species common name.

Description: Body fusiform, elongate, moderately laterally compressed. Dorsally olive, bronze or gold; laterally golden yellow to silver, scales without dark spots at anterior base; ventrally white; dorsal and caudal fins slate colored; anal, pelvic and pectoral fins clear to light orange-red. Head moderately large. Snout blunt. Eye moderately large, placed laterally on upper portion of head. Mouth large, inferior; jaws do not extend backward to anterior end of eye. Lips plicate; posterior margin of lower lip plicae broken into individual papillae; halves of lower lip in a straight line or at an obtuse angle; lower lip thicker than upper lip. Teeth absent on jaws. Pharyngeal teeth comblike, numerous.

Gill rakers 20-32. Dorsal fin rays 12-13, slightly concave distal end. Adipose fin absent.

Caudal peduncle slender, slightly elongate. Circumpeduncular scale rows usually 12.

Caudal fin forked. Anal fin with 7 rays. Pelvic fins with 9 rays; insertion posterior to insertion of dorsal fin. Pectoral fins with 16-19 rays. Lateral line complete with 39-42 large cycloid scales in series. Spawning males develop tubercles on snout and dorsal portion of head, as well as small tubercles on anterior end of body, anal, and caudal fin rays. Spawning females with thickened anal fin rays. Juveniles similar to adults.

Similar Species: Closely resembles other redhorse (Moxostoma) species. Shorthead,

River, and Greater Redhorse all display a red caudal fin and have dark spots present on anterior base of scales on lateral sides. Further, Shorthead Redhorse posterior margin of lower lip forms a straight line. River Redhorse lower lips plicae, but not dissected into 427 individual papillae. Greater Redhorse with 13-15 dorsal fin rays and typically 16 circumpeduncular scale rows. Silver Redhorse have 14-17 rays in the dorsal fin, and halves of lower lip meet at an acute angle.

Distribution and Habitat: Native to east central North America from eastern North and

South Dakota in the north, east throughout the Mississippi, Great Lakes and Hudson Bay drainages to New York, south through the Atlantic Coastal drainages to Alabama and

Mississippi, and west to Oklahoma and Kansas. Limited distribution in the Dakotas with the species primarily occurring within the Red River of the North and Sheyenne river drainages in North Dakota, and well as the Upper Minnesota river drainage in South

Dakota. Inhabit low gradient streams, rivers, lakes, reservoirs and impoundments with warm water temperatures in areas with little aquatic vegetation and coarse substrate.2,8,9

Known to migrate downstream during periods with low water levels.8 Optimum water temperature range reported from Indiana is 26-27.5°C (78.8-81.5°F).6 Rather tolerant of turbidity and high water temperatures, with a suggested upper thermal tolerance of roughly 28.5°C (83.3°F), and a critical thermal maximum of 35.4°C (95.7°F).11

Overwintering habitat consists of deeper water or pools of larger streams.12 Young inhabit areas with low velocity often near stream banks where they seek protection from predators.8

Reproduction: Spawning takes place during spring in water temperatures 10-22.5°C (50-

72.5°F).4,7 Sexual maturity reached at age 3-6.2 Males aggressively defend territories roughly 0.5 m (1.64 ft) in diameter by ramming or butting other nuptial males before and during spawning periods.7,8,10 Females remain in pools or deeper water nearby until one or more males escort a single female into their spawning territory.7 Dominant males, 428 usually one on each side of the female, will vibrate their bodies against the females for 3-

6 seconds initiating egg and sperm release.7 Spawning takes place during daylight over shoals or riffles 30-60 cm (0.98-1.97 ft) deep with gravel and rubble substrate and low to moderate flow.4,7,10 No nest is constructed and no parental care is given.7 Fecundity from the Des Moines River in Iowa ranged from 6,100 eggs from an age-4, 292.1 mm (11.5 in)

TL female, to 25,350 eggs from an age-6, 398.78 mm (15.7 in) TL female.8 Eggs roughly

2.2-2.5 mm (0.09-0.10 in) in diameter and adhesive.1

Age and Growth: Average 97 mm (3.82 in) FL after the first year.10 Largest increases in length occur between ages 2-4, with growth increments decreasing after age-4.10 Females generally larger than males.4 Mean lengths-at-age from Ohio reported as: age-1, 97 mm

(3.82 in) FL; age-2, 126 mm (4.96 in) FL; age-3, 166 mm (6.54 in) FL; age-4, 205 mm

(8.07 in) FL; age-5, 236 mm (9.29 in) FL; age-6, 265 mm (10.43 in) FL; age-7, 290 mm

(11.42 in) FL; age-8, 312 mm (12.28 in) FL.10 Capable of reaching 625 mm (24.6 in)

TL.1,3 Longevity 12 years.2

Food and Feeding: Benthivore. Feeds by sucking material off substrate, and expelling out undesirable prey.10 Foraging most prevalent at night.10 Adults mainly consume immature insects such as Chironomids, Ephemeropterans, and Trichopterans.8,11 Also forages on smaller amounts of filamentous algae and detritus.10,11 Worms and small fingernail clams are also consumed when available.5

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Beckman, D.W., and D.T. Howlett. 2013. Otolith annulus formation and growth

of two redhorse suckers (Moxostoma: Catostomidae). Copeia 2013:390-395. 429

3. Carlander, K.D. 1969. Handbook of freshwater fishery biology, volume 1. Iowa

State University Press, Ames.

4. Curry, K.D., and A. Spacie. 1984. Differential use of stream habitat by spawning

catostomids. The American Midland Naturalist 111:267-279.

5. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

6. Gammon, J.R. 1973. The effects of thermal inputs on the populations of fish and

macroinvertebrates in the Wabash River. Purdue University Water Resources

Research Center Technical Reports, No. 32.

7. Kwak, T.J., and T.M. Skelly. 1992. Spawning habitat, behavior, and morphology

as isolating mechanisms of the golden redhorse, Moxostoma erythrurum, and the

black redhorse, M. duquesnei, two syntopic fishes. Environmental Biology of

Fishes 34:127-137.

8. Meyer, W.H. 1962. Life history of three species of redhorse (Moxostoma) in the

Des Moines River, Iowa. Transactions of the American Fisheries Society 91:412-

419.

9. Page, L.M., and B.M. Burr. 1991. A field guide to freshwater fishes of North

America north of Mexico. Houghton Mifflin, Boston, Massachusetts.

10. Smith, C.A. 1977. The biology of three species of Moxostoma (Pisces-

Catostomidae) in Clear Creek, Hocking, and Fairchild counties, Ohio, with

emphasis on the golden redhorse, M. erythrurum (Rafinesque). Doctoral

dissertation, Ohio State University, Columbus. 430

11. Spiegel, J.R., M.C. Quist, and J.E. Morris. 2011. Trophic ecology and gill raker

morphology of seven catostomid species in Iowa rivers. Journal of Applied

Ichthyology 27:1159-1164.

12. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

431

Shorthead Redhorse, Moxostoma macrolepidotum (Lesueur, 1817)

Etymology and Synonyms: Moxo = Greek for myxo, meaning, “to suck”, stoma = Greek for “mouth”; macrolepidotum = “large scaled”.

Description: Body fusiform, elongate, and moderately laterally compressed. Dorsally olive, bronze or gold; laterally gold to silver; dark crescent shaped spots present at anterior base of scales present; ventrally white; dorsal, anal and paired fins with faint reddish-orange coloration; caudal fin red. Head relatively short and small. Snout moderately blunt; slightly overhanging the mouth. Eyes moderately large, placed laterally on upper portion of head. Mouth small, inferior; jaws do not extend backward to anterior end of eye. Lips plicate with the posterior margin of the lower lip forming a straight line; posterior lower lip plicae dissected into papillae. Teeth absent on jaws. Pharyngeal teeth compressed and comblike with 12-30 on lower half of tooth row. Dorsal fin with 12-14 rays and deeply concave distal end; anterior rays do not extend to end of posterior ray when depressed. Adipose fin absent. Caudal peduncle elongate, moderately thick; circumpeduncular scale rows 12-13. Caudal fin forked. Anal fin with 7 rays. Pelvic fins with 9 rays; insertions posterior to insertion of dorsal fin. Lateral line complete with 41-

45 fairly large cycloid scales in series. Spawning males develop tubercles on the anal and caudal fins and sometimes on the lateral sides. Spawning females develop small tubercles on the head and dorsal surface.

Similar Species: Closely resembles other redhorse (Moxostoma) species. Greater

Redhorse generally have 16 scales in the circumpeduncular row, and have the two halves of the lower lip meeting at an obtuse angle. River Redhorse have the two halves of the lower lip forming a slight obtuse angle, and have a lower lip that is not dissected into 432 papillae. Silver and Golden Redhorse both display a slate colored caudal fin, and generally lack dark spots on the anterior base of scales on the lateral sides.

Distribution and Habitat: Shorthead Redhorse are the most widely distributed species of redhorse in the United States.5,11 Native to the Missouri, Mississippi and Great Lakes-

St. Lawrence basins, and many drainages of the southwestern Hudson Bay basin and the

Atlantic Slope drainages.5,11 Native to all main river basins east and west of the Missouri

River in North Dakota.4 Native to the Red River of the North, James, Sheyenne, Missouri,

Cannonball, Heart, Knife and Little Missouri River basins of North Dakota. Shorthead

Redhorse inhabit clear to slightly turbid waters with increased velocity in large streams with gravel bottoms.8,11 The species also occurs in shallower waters of lakes, reservoirs, and pools of small rivers.1,6,8,11 Shorthead Redhorse appear to seek deeper water during the winter months.11

Reproduction: Spawning migrations begin to occur in April when water temperatures reach roughly 10-15°C (50-59°F), and generally coincide with an increased discharge event.3,9,11 Age at sexual maturity ranges from age-3 to age-5, however the majority of reproductively active males and females are at least age-6.3,11 Spawning activity has been observed to occur during daylight hours in riffle areas with current velocities of 0.3-0.7 m/sec at depths of 20-90 cm (7.87-35.43 in) over fine sand, gravel and cobble substerate.3

The spawning behavior of Shorthead Redhorse consists of one female situated over a riffle area, with 1-6 males positioned either behind or alongside her.3 Rarely do a single male and female spawn together.2,11 The male or males swim up to approach the female, and vibrate or nudge their bodies several times against hers to initiate the release of eggs and milt.3 The average fecundity of females ages 5-7 from Illinois was approximately 433

18,000, which was similar to the average fecundity of females from the Des Moines

River in Iowa.7,11 In Manitoba, 310-418 mm (12.20-16.46 in) FL females contained

12,660-44,329 eggs, with fecundity increasing with the length and age of the female.3

Eggs are >0.60 mm (0.24 in) in diameter, demersal, and adhere to rocks or gravel substrate.3,11

Age and Growth: Females tend to be larger than males.9,11 Mean lengths-at-age from

Iowa are reported as: age-1, 116 mm (4.57 in) TL; age-2, 224 mm (8.82 in) TL; age-3,

293 mm (11.54 in) TL; age-4, 342 mm (13.46 in) TL; age-5, 418 mm (16.46 in) TL; age-

6, 521 mm (20.51 in) TL; age-7, 589 mm (23.20 in) TL; age-8, 655 mm (25.79 in) TL.7

Mean lengths-at-age of individuals from Illinois are reported as: age-1, 93 mm (3.66 in)

TL; age-2, 207 mm (8.15 in) TL; age-3, 295 mm (11.61 in) TL; age-4, 343 mm (13.50 in)

TL; age-5, 368 mm (14.49 in) TL; age-6, 390 mm (15.35 in) TL; age-7, 396 mm (15.60 in) TL; age-8, 402 mm (15.83 in) TL.11 Capable of reaching 615 mm (24.21 in) TL.10

Longevity 20 years.10

Food and Feeding: Young-of-the-year, or individuals ≤ 100 mm (3.94 in) FL feed throughout the water column on smaller zooplankton organisms such as Cladocerans,

Copepods, Ostracods and even some smaller benthic organisms such as Chironomids and

Trichopterans along the bottom.3 Adults, or individuals ˃100 mm (3.94 in) FL consume a wider variety of prey items, and have a predominantly benthic diet made up of

Chironomids, Trichopterans, Ephemeropterans, and mollusks.3 Sand, algae, and other benthic organic material have also been observed in the species diet, as a result of feeding along the bottom.3,11

Literature Cited: 434

1. Becker, G.C. 1963. The fishes of Wisconsin. University of Wisconsin Press,

Madison, Wisconsin.

2. Burr, B.M., and M.A. Morris. 1977. Spawning behavior of the Shorthead

Redhorse, (Moxostoma macrolepidotum) in Big Rock Creek, Illinois.

Transactions of the American Fisheries Society 106:80-82.

3. Harbicht, S. 1990. Ecology of the Shorthead Redhorse (Moxostoma

macrolepidotum, Leseur 1817) in Dauphin Lake, Manitoba. M.S. Thesis,

University of Manitoba.

4. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry, Jr. 2007.

Zoogeographic patterns and faunal change of South Dakota fishes. Western North

American Naturalist 67:161-184.

5. Jenkins, R.E. 1970. Systematic studies of the catostomid fish tribe

Moxostomatini. Part I and Part II. Doctoral dissertation. Cornell University,

Ithaca, New York.

6. Larimore, R.W., and P.W. Smith. 1963. The fishes of Champaign County,

Illinois, as affected by 60 years of stream changes. Illinois Natural History Survey

Bulletin 28:299-382.

7. Meyer, W.H. 1962. Life history of three species of redhorse (Moxostoma) in the

Des Moines River, Iowa. Transactions of the American Fisheries Society 91:412-

419.

8. Pflieger, W.L. 1977. The fishes of Missouri (revised). Missouri Department of

Conservation, Jefferson City, Missouri. 435

9. Reid, S.M. 2006. Timing and demographic characteristics of redhorse spawning

runs in three Great Lakes river basins. Journal of Freshwater Ecology 21:249-258.

10. Reid, S.M. 2007. Comparison of scales, pectoral fin rays and opercles for age

estimation of Ontario redhorse, Moxostoma, species. Canadian Field-Naturalist

12:29-34.

11. Sule, M., and T. Skelly. 1985. The life history of the shorthead redhorse,

Moxostoma macrolepidotum, in the Kankakee River drainage, Illinois. Illinois

Natural History Survey, Biological Notes Number 123:1-16.

436

Greater Redhorse, Moxostoma valenciennesi (Jordan, 1885)

Etymology and Synonyms: Moxo = Greek for myxo, meaning, “to suck”, stoma = Greek for “mouth”; valenciennesi = referring to M.A. Valenciennes, a French naturalist who first described the species.

Description: Body fusiform, elongate, slightly laterally compressed. Dorsally olive, bronze or gold; laterally golden yellow to silver, scales with dark spots present at anterior base; ventrally white; dorsal, caudal and anal fins dark red in color; pelvic and pectoral fins reddish-orange with anterior rays cream in color. Head moderately large. Snout moderately blunt. Eye moderately large, placed laterally on upper portion of head. Mouth large, inferior; jaws do not extend backward to anterior end of eye. Lips plicate with halves of lower lip meeting at an obtuse angle; lower lip thicker than upper lip. Teeth absent on jaws. Pharyngeal teeth thick, numerous. Dorsal fin rays 13-15, slightly convex distal end. Adipose fin absent. Caudal peduncle slender, slightly elongate.

Circumpeduncular scale rows usually 16. Caudal fin forked. Anal fin with 7 rays. Pelvic fins with 9 rays; insertion posterior to insertion of dorsal fin. Pectoral fins with rounded distal end. Lateral line complete with 40-45 large cycloid scales in series. Spawning males develop small tubercles on dorsal and lateral sides of head and body, as well as on rays on lower lobe of caudal fin. Spawning females with small and fewer tubercles present on caudal, anal, pelvic and pectoral fins. Juveniles may have slightly concave dorsal fin.

Similar Species: Closely resembles other redhorse (Moxostoma) species. Silver and

Golden Redhorse both display a slate colored caudal fin, and generally lack dark spots on anterior base of scales on lateral sides. Further, Silver Redhorse posterior margin of lower 437 lip forms an acute angle. Golden Redhorse posterior margin of lower lip forms a nearly straight line or an obtuse angle, and have 39-42 lateral line scales. Shorthead Redhorse with a falcate to concave distal end of dorsal fin and posterior margin of lower lip forms a straight line. River Redhorse with a straight to concave distal end of dorsal fin, and posterior margin of lower lip forms a straight line to slightly obtuse angle.

Distribution and Habitat: Native to the Great Lakes and St. Lawrence River basins, as well as the northern portion of the Ohio, upper Mississippi River, Red River of the North and upper Illinois River basins.3,7 Occurs within the Red and Minnesota river basins of eastern North and South Dakota. Rather rare or uncommon in most of its distribution.1

Inhabit medium to large sized rivers and streams with pool-riffle-run habitat, clear water and sand, gravel, or boulder substrate in areas with slow moving current.2,11 Also known to occupy large lakes and reservoirs.11 Post-spawn and summer home range habitat consists of runs in streams with surface velocities <5 cm/s with cobble or gravel substrate covered with Cladophora.2 Some individuals may also occupy riffle and pool habitat, but to a much lesser extent.2 Migration to overwintering habitat occurs in early fall.2 Sensitive to pollution, siltation and turbidity which likely affects their distribution.1,2

Reproduction: Reproductive ecology of the Greater Redhorse is poorly understood.2,7,8

Spawning migrations consist of movements made upstream in early spring towards riffle areas with increased water velocity and sand, gravel, or cobble substrate.1 Males likely reach sexual maturity at age 5-6.1,5 Spawning is said to occur May-June, however in the

Thousand Island region of the St. Lawrence river, spawning took place from late June through early July in water temperatures 16.7-18.9°C (62.06-66.02°F).1,10 Spawning has been observed in riffle habitat at a mean water depth of 34.4 cm (1.13 ft.) and a mean 438 surface velocity of 38 cm/s.4 Males defend territories which are visited by females periodically to spawn.1 Two males typically spawn with one female.1 No nest is constructed, and no parental care is given. In the Grand River in Ontario, Canada, spawning ends early June, which is noted to be earlier than other larger and cooler rivers.2,6,8 After spawning ceases, adults move as far as 15.2 km (9.44 mi) back downstream.2 A spike in water temperature or heavy rain event likely leads to the termination of seasonal spawning.2 Eggs yellow, roughly 3.0-3.5 mm (0.12-0.14 in) in diameter, nonadhesive, demersal and settle into spaces between substrate.3 Artificially spawned eggs hatched within 6-8 days at 16-19°C (60.8-66.2°F).3

Age and Growth: Largest of the redhorse species.1 Little information available on growth rates. Growth rates reported from Wisconsin vary with population and distribution.1 Newly hatched larvae 9-14 mm (0.35-0.55 in) TL.3 Females often larger than males.2 Capable of reaching 688 mm (27.09 in) TL.9 Longevity 20 years.9

Food and Feeding: Benthivore. Little information available on diet and feeding ecology.

Like most castostomid species, Greater Redhorse feed by sucking material off substrate, and expelling out undesirable prey. Adults mainly consume aquatic benthic invertebrates such as midge larvae, crustaceans, and mollusks.1

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Bunt, C.M., and S.J. Cooke. 2001. Post-spawn movements and habitat use by

greater redhorse, Moxostoma valenciennesi. Ecology of Freshwater Fish 10:57-

60. 439

3. Bunt, C.M., and S.J. Cooke. 2004. Ontogeny of larval greater redhorse

(Moxostoma valenciennesi). The American Midland Naturalist 151:93-100.

4. Cooke, S.J., and C.M. Bunt. 1999. Spawning and reproductive biology of greater

redhorse, Moxostoma valenciennesi, in the Grand River, Ontario. Canadian Field-

Naturalist 113:497-502.

5. Jenkins, R.E. 1970. Systematic studies of the catostomid fish tribe

Moxostomatini. Doctoral dissertation, University of Michigan, Ann Arbor.

6. Jenkins, R.E., and D.J. Jenkins. 1980. Reproductive behavior of the greater

redhorse, Moxostoma valenciennesi, in the Thousand Islands Region. Canadian

Field-Naturalist 94:426-430.

7. Kay, L.K., R. Wallus, and B.L. Yeager. 1994. Reproductive biology and early life

history of fishes in the Ohio River drainage. Volume 2: Catostomidae.

Chattanooga: Tennessee Valley Authority.

8. Mongeau, J.R., P. Dumont, and L. Cloutier. 1992. La biologie du suceur cuivré

(Moxostoma hubbsi) compare á celle de quatre autres espèces de Moxostoma (M.

anisurum, M. carinatum, M. macrolepidotum et M. valenciennesi). Canadian

Journal of Zoology 70:1354-1363.

9. Reid, S.M. 2007. Comparison of scales, pectoral fin rays, and opercles for age

estimation of Ontario Redhorse, Moxostoma, species. Canadian Field-Naturalist

121:29-34.

10. Scott, W.B. 1967. Freshwater fishes of eastern Canada. University of Toronto

Press, Toronto. 440

11. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

441

CHAPTER 12

FAMILY ICTALURIDAE

Introduction

Around the world, there are roughly 34 families and well over 2,000 species of marine and freshwater catfishes. The North American Catfish family, Ictaluridae, is made of 46 species and is the only catfish family that occurs in North America. The native distribution of Ictaluridae includes the freshwaters essentially from the Continental

Divide east to the Atlantic Coast, but they have been widely introduced outside their native range throughout North America. Ictalurids can easily be identified from other families of fishes in the Dakotas by their scaleless body, a present flap-like adipose fin

(sometimes connected to the caudal fin), 4 sets of barbels, and spines present in the pectoral fins. Barbels have numerous tiny tasebuds located in them, which assist

Ictalurids in locating prey. Pectoral and dorsal spines in most Ictalurids, including the more popular sportfish, Channel Catfish, Ictalurus punctatus, are capable of injecting a small amount of venom, creating a painful sting that may be intensely painful, but non- life threatening. Species in the family Ictaluridae also range in a wide variety of lengths.

Many people often associate the family with the larger catfish species like the Channel

Catfish, the Flathead Catifish, Pylodictis olivaris, the Blue Catfish, Ictalurus furcatus, and even the more common bullheads (Ameiurus). However, the family also includes numerous smaller species including the Tadpole Madtom, Noturus gyrinus, and the

Stonecat, Noturus flavus, (the only two species of Noturus that occur in the Dakotas) which are generally less than 250mm (10in) in total length. 442

The larger members of the family that occur within the Dakotas inhabit large rivers and main tributaries, with the exception of the Channel Catfish that are occasionally found in larger creeks. Bullheads primarily inhabit lakes, ponds and streams and are highly tolerant of turbidity and low dissolved oxygen levels. Madtoms prefer small streams, sloughs, ponds and lakes with silt, sand, pebble or gravel substrates.

Ictalurids are generally most active during the night when they migrate from deep to shallow waters to forage during dark. The family is well adapted to forage during the night given the numerous taste buds located on the body and barbels, which makes up for their lack of visual enhancements. All catfish are considered opportunistic nocturnal predators, and undergo an ontogenetic diet shift to piscivory as they reach adulthood.

Bullheads are also opportunistic nocturnal foragers, but adults consume a wider variety of prey items including zooplankton, benthic aquatic invertebrates, mollusks and small fish.

Madtoms are also nocturnal and primarily feed along the bottom substrate on organic debris, zooplankton and aquatic insect larvae. Males, and sometimes females of certain species, construct nests, or depressions within the substrate or cavities within submerged structures, where they provide parental care by fanning the eggs until they hatch.

Bullheads are even known to guard and swim alongside their young, which form a tight school often called a “ball”.

Channel Catfish, Flathead Catfish, and even the less frequent Blue Catfish are all important sportfish species across the Dakotas, whereas bullheads and madtoms often serve as forage species for other important sportfish such as Walleye, Sander vitreus, and

Northern Pike, Esox lucius.

443

Black Bullhead, Ameiurus melas (Rafinesque, 1820)

Etymology and Synonyms: Ameiurus = Greek, a- = “without”, -meiouros = “without tail”, referring to the unforked caudal fin; melas = “black”, referring to the species coloration and common name.

Description: Body robust, stout, rounded anteriorly; laterally compressed posteriorly.

Dorsally solid black, dark brown, or dark olive green; laterally gray, tan-brown; ventrally yellow-cream to white; faint yellow-cream vertical bar at base of caudal fin; fins uniformly dark in color; all barbels dark gray, brown, to black. Head slightly dorsoventrally flattened. Snout blunt. Eye small, positioned dorsolaterally on head.

Mouth terminal, wide; jaws nearly equal in length with upper jaw occasionally slightly extending past lower jaw. Four pairs of barbels total; one pair on top of snout; one long pair near corners of mouth on upper jaw; two pairs on chin placed in a transverse line.

Tooth patches present on both jaws with numerous sharp, small teeth. Gill rakers 15-19.

Dorsal fin with single stout spine and 5-6 rays. Adipose fin present, separated from caudal fin, flap-like. Caudal peduncle slender, short. Caudal fin weakly forked, almost square in shape. Anal fin elongate with rounded distal end, 16-21 rays. Pelvic fin small, abdominal with 7-8 rays. Pectoral fin with single, stout spine weakly serrated on posterior edge. Lateral line complete. Scaleless. Juveniles and young adults much darker in color.

Similar Species: Closely resembles the Yellow and Brown Bullhead. Yellow Bullhead often lighter in color with no vertical bar at base of caudal fin, cream to white barbels on chin, 24-27 anal fin rays, and 12-16 gill rakers. Brown Bullhead often with brown-black mottling on dorsal and lateral sides, (although occasionally a solid color), 20-24 anal fin rays, and 12-15 gill rakers. Flathead Catfish with a more pronounced dorsoventrally 444 flattened head with a lower jaw extending past the upper jaw. Stonecat and Tadpole

Madtom have an adipose fin strongly fused with the caudal fin.

Distribution and Habitat: Native throughout majority of central North America east of the Rocky Mountains, from extreme eastern Montana, through southern Canada, east to eastern New York and Pennsylvania, and south, west of the Appalachians to the Gulf of

Mexico. Widely introduced outside its native range. Abundant and widespread throughout the Dakotas. Occurs in a wide variety of habitats including wetlands, ponds, lakes, reservoirs, streams and rivers. In lotic habitats, the species is often found in low gradient areas with low velocity such backwaters or pools with dense cover or aquatic vegetation. Often found over gravel, silt, or mud substrates. In eastern South Dakota lakes, abundance generally decreases with increasing lake area, volume, and depth.2 Most frequent in small, shallow, turbid lakes with plentiful shoreline vegetation.2 Tolerant of a wide range in turbidities, high water temperature, and low dissolved oxygen. Adults rather sensitive to daylight and typically inactive during the day, hiding under vegetated cover; most active at night. Juvenile habitat requirements similar to adults.8 Upper lethal limit for all life stages is 35-39°C (95-102°F).8

Reproduction: Spawning takes place mid-May to early July, or when water temperature reaches 20°C (68°F).3,7,8 Age at sexual maturity variable, but typically around 160 mm

(6.29 in) TL. Females known to construct saucer-shaped nests roughly 15-35 cm (5.9-

13.8 in) in diameter using her snout, pelvic, and anal fins.5,10 Nest are created in areas 0.6-

1.2 m (2-4 ft.) in depth, near vegetative cover or woody debris in sand, silt, or mud substrates.10 Eggs yellow, roughly 0.8-1.6 mm (0.03-0.06 in) in diameter.1 Following egg fertilization, parental care is given by both male and female by fanning the gelatinous egg 445 masses. Hatching occurs within 5-10 days depending on water temperature. Fry known to form tight schools, or ball, near the surface guarded by the adults for two weeks or longer.1

Age and Growth: Growth highly variable among populations and density dependent with more abundant populations experiencing slower growth.2,4 In South Dakota, individuals known to grow larger in deeper lakes than from shallow lakes with a larger population.4 Stunting is also known to occur in dense populations with less predators.4 In the James River, growth known to be faster in years with high spring flows.9 Although survival rates likely increase in turbid water, growth rates increase in clear waters.4 Mean lengths-at-age from eastern South Dakota lakes are reported as: age-1, 101 mm (3.98 in)

TL; age-2, 155 mm (6.10 in) TL; age-3, 203 mm (7.99 in) TL; age-4, 232 mm (9.13 in)

TL; age-5, 262 mm (10.31 in) TL; age-6, 289 mm (11.37 in) TL.4 Adults average <305 mm (12 in) TL, but capable of exceeding 381 mm (15 in) TL. Longevity 10 years.

Food and Feeding: Opportunistic, primarily selective zooplanktivores. Diet does not vary greatly with size, and all stages known to be selective of the largest cladocerans possible.6 Young-of-year primarily prey upon limnetic cladocerans and copepods.6

Adults primarily feed on zooplankton, but also scavenge on insect larvae, crayfish, snails, and small fish. Majority of feeding takes place near the bottom at night in shallow waters.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Brown, M.L., D.W. Willis, and B.G. Blackwell. 1999. Physiochemical and

biological influences on black bullhead populations in eastern South Dakota

glacial lakes. Journal of Freshwater Ecology 14(1):47-60. 446

3. Dennison, S.G., and R.V. Bulkley. 1972. Reproductive potential of the black

bullhead, Ictalurus melas, in Clear Lake, Iowa. Transactions of the American

Fisheries Society 101:483-487.

4. Hanchin, P.A., D.W. Willis, and M.J. Hubers. 2002. Black bullhead growth in

South Dakota waters: limnological and community influences. Journal of

Freshwater Ecology 17: 65-73.

5. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

6. Repsys, A.J. 1972. Food selectivity of the Black Bullhead (Ictalurus melas,

Rafinesque) in Lake Poinsett, South Dakota. M.S. thesis, South Dakota State

University, Brookings.

7. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

8. Stuber, R.J. 1982. Habitat suitability index models: Black bullhead. U.S. Fish and

Wildlife Service, FWS/OBS-82/10.14, Washington, DC.

9. Tol, D. 1976. An evaluation of the fishery resources in a portion of the James

River, South Dakota scheduled for channel modification. M.S. thesis, South

Dakota State University, Brookings.

10. Wallace, C.R. 1967. Observations on the reproductive behavior of the black

bullhead (Ictalurus melas). Copeia 1967(4):852-53.

447

Yellow Bullhead, Ameriurus natalis (Lesueur, 1819)

Etymology and Synonyms: Ameiurus = Greek, a- = “without”, -meiouros = “without tail”, referring to the unforked caudal fin; natalis = Latin for “having a large buttocks”.

Description: Body robust, stout, rounded anteriorly; laterally compressed posteriorly.

Dorsally solid yellow-olive to dark brown-black in color; laterally yellow-olive to brown; ventrally yellow-cream to white; light vertical bar at base of caudal fin absent; fins dusky dark olive in color; barbels on chin cream to white. Head slightly dorsoventrally flattened. Snout blunt. Eye small, positioned dorsolaterally on head. Mouth terminal, wide; jaws nearly equal in length with upper jaw occasionally slightly extending past lower jaw. Four pairs of barbels present; one pair on top of snout; one long pair near corners of mouth on upper jaw; two pairs on chin placed in a transverse line. Tooth patches present on both jaws with numerous sharp, small teeth. Gill rakers 12-16. Dorsal fin with single stout spine and 5-6 rays. Adipose fin present, separated from caudal fin, flap-like. Caudal peduncle slender, short. Caudal fin weakly forked, almost square in shape. Anal fin elongate with nearly straight distal end, 24-27 rays. Pelvic fin small, abdominal with 7-8 rays. Pectoral fin with single, stout spine distinctly serrated on posterior edge. Lateral line complete. Scaleless. Juveniles and young adults with less yellow coloring.

Similar Species: Closely resembles the Black and Brown Bullhead. Black Bullhead with dark gray, brown, to black barbels on chin, 16-21 anal fin rays, and 15-19 gill rakers.

Brown Bullhead often with brown-black mottling on dorsal and lateral sides, (although occasionally a solid color), 20-24 anal fin rays, and 12-15 gill rakers. Flathead Catfish with a more pronounced dorsoventrally flattened head with a lower jaw extending past 448 the upper jaw. Stonecat and Tadpole Madtom have an adipose fin strongly fused with the caudal fin.

Distribution and Habitat: Native throughout the eastern United States, west into southeastern Canada and the Great Plains, and South to the Gulf of Mexico. Less abundant in the Dakotas than the Black Bullhead; occurs throughout the Missouri River and its major tributaries. Occurs in a wide variety of habitats including wetlands, ponds, lakes, reservoirs, streams and rivers. Most common in shallow, low flow areas of lakes and streams with clear water and abundant aquatic vegetation. Often occurs at depths of

0.5-1.5 m (1.64-4.92 ft.) over gravel, sand, or mud substrates. Periodically found in slightly turbid to turbid waters. Tolerant of high water temperatures, and low dissolved oxygen levels.

Reproduction: Spawning generally earlier than Black or Brown Bullhead, taking place

May to July. Age at sexual maturity variable, but typically at age 2-3, or around 140 mm

(5.51 in) TL.3 Spawning ritual similar to the Black Bullhead and consists of the male and female laying side-by-side in opposing directions, vibrating their bodies together.4 One or both adults known to construct saucer-shaped nests, often located near submerged structure. Also known to spawn in natural cavities within rocks or submerged trees. Eggs adhesive, yellow-cream in color, and roughly 2.2-3.0 mm (0.08-0.12 in) in diameter.2

Following egg fertilization, parental care is given by one or both adults by fanning the gelatinous egg masses. Fecundity 1,650-4,300 eggs per female.2 Roughly 300-700 eggs are laid per nest.2 Hatching occurs within 5-10 days depending on water temperature.

Several hundred fry known to form tight schools near the surface guarded by an adult, usually the male, until fry reach 50 mm (1.97 in) TL.2 449

Age and Growth: Information on growth is scarce. Generally smaller than the Black or

Brown Bullhead. Growth highly variable among environmental conditions and populations. Mean length-at-age from Wisconsin was reported as: age-2, 206 mm (8.11 in) TL; age-3, 226 mm (8.89 in) TL; age-4, 272 mm (10.71 in) TL.2 Capable of reaching

298 mm (11.73 in) TL.1 Longevity 12 years.1

Food and Feeding: Opportunistic, voracious omnivore. Prey items similar to the Black

Bullhead, however a greater amount of aquatic vegetation and bottom ooze is generally consumed. Able to locate prey with taste buds present all over the body, especially on the barbels. Adults consume a wide variety of prey items including small fish, benthic invertebrates, mollusks, and macrophytes.5 Majority of feeding takes place near the bottom at night in shallow waters.5 Adults known to scavenge on dead fish after winterkill.6

Literature Cited:

1. Murie, D.J., D.C. Parkyn, W.F. Loftus, and L.G. Nico. 2009. Variable growth and

longevity of yellow bullhead (Ameiurus natalis) in the Everglades of south

Florida, USA. Journal of Applied Ichthyology 25:740-745.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

4. Wallace, C.R. 1967. Observations on the reproductive behavior of the black

bullhead (Ictalurus melas). Copeia 1967(4):852-53.

5. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 450

6. Schneider, J.C. 1998. Fate of dead fish in a small lake. American Midland

Naturalist 140:192-196.

451

Brown Bullhead, Ameriurus nebulosus (Lesueur, 1819)

Etymology and Synonyms: Ameiurus = Greek, a- = “without”, -meiouros = “without tail”, referring to the unforked caudal fin; nebulosus = “clouded” referring to the species mottled coloring.

Description: Body robust, stout, rounded anteriorly; laterally compressed posteriorly.

Dorsally mottled dark brown to black (although occasionally a solid color); laterally mottled with light and dark brown; ventrally cream to white; light vertical bar at base of caudal fin absent; fins dark in color, may be mottled; barbels black to yellow-brown.

Head slightly dorsoventrally flattened. Snout blunt. Eye small, positioned dorsolaterally on head. Mouth terminal, wide; jaws nearly equal in length with upper jaw occasionally slightly extending past lower jaw. Barbels present, four pairs total; one pair on top of snout; one long pair near corners of mouth on upper jaw; two pairs on chin placed in a transverse line. Tooth patches present on upper and lower jaws with numerous sharp, small teeth. Gill rakers 12-15. Dorsal fin with single stout spine and 6-7 rays. Adipose fin present, separated from caudal fin, flap-like. Caudal peduncle slender, short. Caudal fin weakly forked, almost square in shape. Anal fin elongate with rounded distal end, 20-24 rays. Pelvic fin small, abdominal with 7-8 rays. Pectoral fin with single spine distinctly serrated on posterior edge. Lateral line complete. Scaleless. Juveniles and young adults similar to adults, but more likely to be solid in color.

Similar Species: Closely resembles the Black and Yellow Bullhead. Black Bullhead solid black, dark brown to olive green in color, with 16-21 anal fin rays, and 15-19 gill rakers. Yellow Bullhead often lighter in color, also with no vertical bar at base of caudal fin, cream to white barbels on chin, 24-27 anal fin rays, and 12-16 gill rakers. Flathead 452

Catfish with a more pronounced dorsoventrally flattened head with a lower jaw extending past the upper jaw. Stonecat and Tadpole Madtom have an adipose fin strongly fused with the caudal fin.

Distribution and Habitat: Native throughout southeastern Canada, south to eastern

North and South Dakota, throughout the Mississippi and Hudson Bay river drainages to the Atlantic Coast, and south to Florida. Less abundant in the Dakotas than the Black

Bullhead. Inhabits lakes, reservoirs, rivers, streams, sloughs, and ponds. Most common in low flow, shallow areas of sloughs and ponds with dense aquatic vegetation and gravel, sand, or silt substrates. Tolerant of a wide range in turbidities, high water temperature, and dissolved oxygen. Prefers warm water. Optimal water temperature is 26°C (78.8°F).7

Upper lethal limit reported as 37.5°C (99.5°F).1,5

Reproduction: Spawning takes place mid-May through July, or when water temperature reaches 21-25°C (69.8-77°F).11 Sexual maturity occurs at age 2-3. Adults migrate at night from deep to shallow waters into spawning sites.3 Males court females within nesting areas with nipping and budging behavior.3 Females, and occasionally males, construct saucer-shaped nests often located near submerged structure or vegetation within sand or gravel substrates.1,3 Nests are excavated in waters less than 1 m (39.37 in) deep near the shoreline, and are guarded by males who drive other bullheads away.1,3 Spawning ritual similar to Black and Yellow Bullhead. Spawning takes place early morning to midafternoon, and consists of the male and female laying side-by-side in opposing directions vibrating their bodies together.1,4 Fecundity increases with size of female.

Individuals 202-230 mm (7.95-9.05 in) TL produced roughly 2,000-13,000 eggs annually.8 Eggs adhesive, cream in color, and roughly 3.0 mm in diameter.1,4 One or both 453 adults give parental care by fanning the gelatinous egg masses with the caudal, anal, and pectoral fins.3 Hatching occurs within 6-9 days at water temperatures of 20.6-23.3°C

(69.1-73.9°F), and may take up to 13 days.1,3 After hatching, fry remain in the nest for 7-

10 days, after which they begin to swim and form tight schools guarded by the parents from potential predators such as sunfish, bass, and juvenile perch.1,2 Parental care is given until fry reach roughly 50 mm (1.97 in) TL.8

Age and Growth: Little information available on growth. Larvae roughly 4-8 mm (0.16-

0.31 in) TL at hatching.11 Growth rates highly variable among environmental conditions and populations. Maximum growth rates occur at 20-25°C (68-77°F).9 Capable of reaching 508-532 mm (20-21 in) TL.6 Longevity 11 years.

Food and Feeding: Opportunistic omnivore. Able to locate prey with taste buds present all over the body, especially on the barbels. Fry and young-of-year primarily consume zooplankton and chironomids.6 Adults forage on a variety of prey items including benthic invertebrates, small fish, aquatic vegetation, and mollusks. Feeding takes place both day and night, primarily along the bottom. Metabolic rate increases with water temperature.10

Maximum consumption increases with temperature from 10-25°C (50-77°F), and begins declining at 30°C (86°F).9 Consumption known to cease by 35°C (95°F).9

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Blumer, L.S. 1982. Parental care and reproductive ecology of the North American

catfish, Ictalurus nebulosus. Ph.D. Dissertation, University of Michigan, Ann

Arbor. 454

3. Blumer, L.S. 1985. Reproductive natural history of the Brown Bullhead, Ictalurus

nebulosus in Michigan. The American Midland Naturalist 114(2):318-330.

4. Breder, C.M., Jr., and D.E. Rosen. 1966. Modes of reproduction in fishes.

American Museum of Natural History, New York. 941pp.

5. Brett, J.R. 1944. Some lethal temperature relations of Algonquin Park fishes.

University of Toronto Stud. Biol. Ser. No. 52. 49pp.

6. Carlander, K.D. 1969. Handbook of freshwater fishery biology, volume 1. Iowa

State University Press, Ames.

7. Crawshaw, L.I., and H.T. Hammel. 1974. Behavioral regulation of internal

temperature in the brown bullhead, Ictalurus nebulosus. Comparative

Biochemistry and Physiology Part A: Physiology 47(1):51-60.

8. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

9. Hartman, K.J. 2017. Bioenergetics of brown bullhead in a changing climate.

Transactions of the American Fisheries Society 146:634-644.

10. Keast, A. 1984. Growth responses of the brown bullhead (Ictalurus nebulosus) to

temperature. Canadian Journal of Zoology 63:1510-1515.

11. Mansueti, A.J., and J.D. Hardy. 1967. Development of fishes of the Chesapeake

Bay region. University of Maryland, Baltimore, Natural Resources Institute, Part

I. 202 pp.

455

Blue Catfish, Ictalurus furcatus (Lesueur, 1840)

Etymology and Synonyms: Ictalurus =Greek for “fishcat” -ichtys = fish, Greek, ailouros

= cat; furcatus = Latin for the word “forked” in reference to the tail.

Description: Body deep and robust. Dorsally and laterally blue-silver in color; ventrally white. Spots absent. Larger individuals appear pale blue laterally and gray dorsally.

Juveniles and younger individuals appear silver-white in color. Head broad, wedge- shaped; virtually straight profile from tip of snout to anterior part of dorsal fin. Upper jaw extends further than lower jaw. Barbels present, white in color on chin. Teeth present on inner margin of upper jaw. Dorsal fin with 1 spine anteriorly followed by 6 rays. Adipose fin present. Caudal peduncle concise. Caudal fin deeply forked, separate from adipose fin. Anal fin long and nearly straight with 30-35 rays. Pelvic fin with 8 rays. Pectoral fins with 1 spine and 9 rays. Lateral line complete. Scales absent. Swim bladder with two chambers. Sexual dimorphism shown in the genetial orifices; males with circular opening with prominent papillae; females with slit-like opening and less prominent papillae.2,9

Similar Species: Similar to larger Channel Catfish (Ictalurus punctatus) which can display a bluish hue and lose their spots when breeding. Anal fin of Channel Catfish is rounded, unlike the straight edge of Blue Catfish. Channel Catfish also have a rounder profile from tip of snout to anterior part of dorsal fin, unlike Blue Catfish which have an almost straight profile. Channel Catfish have a single chamber in the swim bladder.

Distribution and Habitat: Native to the Mississippi, Missouri, and Ohio River basins.

Also found in Gulf Coast streams from Alabama to Mexico, and northern Central

America.1,2 Considered native of South Dakota with restricted range. Primarily found in the Missouri River and the lowermost sections of its larger tributaries in South Dakota.3 456

Not reported in North Dakota.2 Primarily considered a big-river species. Occurs in large flowing riverine habitats with deep, swift channels or flowing pools with normal turbidity and a mud or silt substrate.2 Also found in open waters of large reservoirs. Large individuals often reside in tailwaters below dams.2,5 Able to withstand moderately high levels of turbidity.2 Will move into tributaries and backwaters during high water periods.4

Migrates up to several hundred km upstream in the spring and downstream in the fall in response to water temperature.2,6

Reproduction: Spawning time and habits are relatively unknown, but are probably similar to the Channel Catfish (Ictalururs punctatus).6,7,8 Spawning occurs in late spring or early summer within protected areas where currents are minimal. Larger individuals often migrate both upstream and downstream to seek spawning sites.2 Maturity for both males and females is typically reached at an earlier age in the southern portion of their range.2 In Lake of the Ozarks, Missouri, sexual maturity was reached at 42 – 48cm (16.5

– 18.9in) TL and ages 6 – 7.10 Eggs roughly 2.5mm (0.098in) in diameter.2 Mature ova from a 7-9kg (15.43 – 19.84 lbs) female were found to be 3.0-3.3mm (0.001 – 0.010in) in diameter.2 Eggs hatch within 7-8 days in water temperatures of 21 – 24°C (69.8 –

75.2°F).2,6,11 Like other ictalurid catfish species, males are known to guard the eggs and fry.

Age and Growth: Largest catfish in the United States. Growth is rapid, especially after becoming picivorous.2 Growth similar between both sexes. Adults typically >122cm (48 in), but capable of reaching 165cm (65 in) and 57kg (125 lbs.) in the Midwest area.13

Growth rates are known to be faster in southern regions due to an increased growing 457 season, warmer water temperatures, and a more diverse forage base.2 Capable of living 20

– 25 years. 2,12

Food and Feeding: Omnivorous and opportunistic feeders. Smaller individuals consume mostly small invertebrates and some small fish. Larger individuals primarily eat fish and larger invertebrates.6 Blue catfish are also known to feed on fingernail clams and freshwater mussels.2 Feeding takes place along the bottom substrate using their sense of taste and smell more than sight in locating food.2,6

Literature Cited:

1. Glodek, G.S. 1980. Ictalurus furcatus (LeSueur) blue catfish. Page 439 in D.S.

lee, et al. Atlas of North American freshwater fishes. North Carolina State

Museum of Natural History, Raleigh.

2. Graham, K. 1999. A review of the biology and management of blue catfish. In

E.R. Irwin, W.A. Hubert, & C.F. Rabeni (Eds.), Catfish 2000: proceedings of the

international ictalurid symposium. American Fisheries Society Symposium

24:37-49.

3. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

4. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

5. Mettee, M.F., P.E. O’Neil, and J.M. Pierson. 1996. Fishes of Alabama and the

mobile basin. Oxmoor House, Inc., Birmingham, Alabama.

6. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 458

7. Lagler, K.F. 1961. Freshwater fishery biology. William C. Brown Company,

Dubuque, Iowa.

8. Harlan, J.R., E.B. Speaker, and J. Mayhew. 1987. Iowa fish and fishing, fifth

edition. Iowa Department of Natural Resources, DesMoines.

9. Moyle, P.B. 1976. Inland fishes of California, University of California Press,

Berkeley and Los Angeles.

10. Graham, K., and K. DeiSanti. 1999. The population and fishery of blue catfish

and channel catfish in the Harry S. Truman Dam tailwater, Missouri. Pages 361-

376 in Irwin et al. (1999).

11. Henderson, G.G. 1972. Rio Grande blue catfish study. Texas Parks and Wildlife

Department, Federal Aid in Fisheries Restoration Project F-18-R-7, Job 11,

Progress Report, Austin.

12. Hale, R.S., and T.J. Timmons. 1990. Growth of blue catfish in the lacustrine and

riverine areas of the Tennessee portion of Kentucky Lake. Journal of the

Tennessee Academy of Science 65(3):86-90.

13. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

459

Channel Catfish, Ictalurus punctatus (Rafinesque, 1818)

Etymology and Synonyms: Ictalurus = “Fish cat” referring to the barbels resembling a cats whiskers; punctatus = “spotted”, referring to the lateral spots present on juveniles and young adults.

Description: Body slender, elongate, laterally compressed posteriorly. Dorsally dusky blue or pale to slate olive-gray; laterally slate to light gray; ventrally white; fins lightly pigmented; maxillary barbels dark brown, ventral barbels lightly pigmented. Head small, slightly dorsoventrally flattened; profile slightly rounded from tip of snout to anterior end of dorsal fin. Snout blunt. Eye small, positioned dorsolaterally on head. Mouth wide, subterminal; upper jaw extends past lower jaw. Barbels present, four pairs total; one pair on top of snout; one long pair near corners of mouth on upper jaw; two pairs on chin with inner pair shorter than outer pair. Teeth present on upper and lower jaws, small and sharp; premaxillary tooth patch present on upper jaw without backwards extensions. Gill rakers 14-18. Dorsal fin with 1 short spines, 6-7 rays. Adipose fin present, flap-like.

Caudal peduncle slender, short. Caudal fin deeply forked. Anal fin elongate with rounded distal end, 24-29 rays. Pelvic fin small, abdominal with 8-9 rays. Pectoral fin with 1 strongly serrated spine on posterior edge with 7-9 rays. Lateral line complete. Scaleless.

Juveniles and young adults with small, distinct scattered dark spots on lateral sides.

Spawning males more blue-green dorsally.

Similar Species: Closely resembles Blue Catfish. Blue Catfish display an anal fin with

30-35 rays with a nearly straight distal end, and have a straight profile from the tip of the snout to anterior end of dorsal fin. Flathead Catfish have a slightly forked caudal fin, 460 almost square in shape, and a more pronounced dorsoventrally flattened head with a lower jaw extending past the upper jaw.

Distribution and Habitat: Native to southern Quebec and Manitoba, Canada in the north, south throughout the Missouri and Mississippi River basins from Montana in the west, east throughout the St. Lawrence-Great lakes and Hudson Bay drainages to eastern

New York, and south to northern Mexico and the Gulf of Mexico. Habitat generalists; inhabit a wide variety of habitat types including rivers, streams, lake, reservoirs, and ponds. Thrive in low-gradient, well-oxygenated, medium to large turbid rivers with alternating pool and riffle habitat. Often found over mud, sand, gravel, or silt substrates.

During daylight hours, the species prefers dark, deeper pools and crevices under rocks or submerged woody debris or areas with moderate amounts of aquatic vegetation. Juvenile and young individuals inhabit shallow shoals and sand bars near riffles.

Reproduction: Migrations from overwintering habitat to upstream spawning sites begin in the spring.11 Spawning takes place in late spring and early summer when water temperatures reach 22-28°C (71.6-82.4°F). Sexual maturity correlated with size, which depends on length of growing season and location. Northern populations reach sexual maturity at greater lengths than southern populations. Males construct a nest often near undercut stream banks or crevices under rocks and trees. Eggs light yellow, demersal, adhesive, and roughly 3.2 mm (0.13 in) in diameter.2 Fecundity roughly 8,800 eggs/kg of body weight from females 0.45-1.81 kg (1-4 lbs).2,3 Following fertilization of eggs, the male chases the female out of the nesting area, which he will continue to fan and defend until that larvae hatch.10 Hatching occurs within 7-8 days. 461

Age and Growth: Larvae roughly 6.4 mm (0.25 in) TL post hatching.2 Growth rates variable among populations, and may be influenced by localized environmental conditions, climate and resources availability.7,8,13 Low-flow conditions contribute to young channel catfish growth by creating a greater amount of shallow, warm, and low velocity habitats.6 Mean lengths-at-age from the upper segment of the Big Sioux River were reported as: age-2, 216 mm (8.5 in) TL; age-3, 307 mm (12.1 in) TL; age-4, 391 mm (15.4 in) TL; age-5, 456 mm (18.0 in) TL; age-6, 495 mm (19.5 in) TL; age-7, 623 mm (24.5 in) TL; age-8, 631 mm (24.8 in) TL; age-9, 678 mm (26.7 in) TL; age-10, 672 mm (26.5 in) TL.7 Capable of exceeding 1,000 mm (39 in) TL.12 Average longevity 10-

15 years, capable of reaching 27 years.12

Food and Feeding: Adults generalist, opportunistic omnivores with a diverse and variable diet, allowing them to adjust to changing conditions throughout growing seasons and years.13 Migrate from deep to shallow waters near rifles at night to feed. Larvae primarily feed on zooplankton. Juveniles consume macroinvertebrates, and shift to larger macroinvertebrates as they grow.1,13 Undergo and ontogenetic diet shift to piscivory at approximately 280 mm (11 in) TL, however benthic macroinvertebrates remain in the diet of individuals of all sizes.5,13 Invertebrate availability known to influence Channel

Catfish growth and condition.4,11 Adults primarily consume small fish, but also prey upon

Dipterans, annelids, crayfish, mollusks, immature Ephemeroptera and Trichoptera, neamatodes, and filamentous algae.9,13 Also known to occasionally consume birds and rodents.

Literature Cited:

462

1. Bailey, R.M., and H.M. Harrison. 1948. Food habits of the southern channel

catfish (Ictalurus Lacustris punctatus) in the Des Moines River, Iowa.

Transactions of the American Fisheries Society 75(1):110-138.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Clemens, H.P., and K.E. Sneed. 1957. The spawning behavior of the channel

catfish Ictalurus punctatus. U.S. Fish and Wildlife Service Special Scientific

Report. Fish. No. 219. 11pp.

4. Hampton, D.R., and C.R. Berry, Jr. 1997. Fishes of mainstem Cheyenne River in

western South Dakota. Proceedings of the South Dakota Academy of Science

76:11-25.

5. Hill, T.D., W.G. Duffy, and M.R. Thompson. 1995. Food habits of channel

catfish in Lake Oahe, South Dakota. Journal of Freshwater Ecology 10:319-323.

6. Hogberg, N.P., M.J. Hamel, and M.A. Pegg. 2016. Age-0 channel catfish

Ictalurus punctatus growth related to environmental conditions in the channelized

Missouri River, Nebraska. River Research and Applications 32:744-752.

7. Kirby, D.J. 2001. An assessment of the channel catfish population in the Big

Sioux River, South Dakota. M.S. Thesis, South Dakota State University,

Brookings.

8. Pegg, M.A., and C.L. Pierce. 2001. Growth rate responses of Missouri and Lower

Yellowstone river fishes to a longitudinal gradient. Journal of Fish Biology

59:1529-1543. 463

9. Peterson, E. 2017. Invertebrate prey selectivity of channel catfish (Ictalurus

punctatus) in western South Dakota prairie streams. M.S. Thesis, South Dakota

State University, Brookings.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

11. Quist, M.C., and C.S. Guy. 1998. Population characteristics of channel catfish

from the Kansas River, Kansas. Journal of Freshwater Ecology 13:351-359.

12. Siddons, S.F., and M.A. Pegg. 2016. Age, growth, and mortality of a trophy

Channel Catfish population in Manitoba, Canada. North American Journal of

Fisheries Management 36:1368-1374.

13. Stevens, T.M. 2013. Feeding ecology and factors influencing growth and

recruitment of channel catfish in South Dakota reservoirs. M.S. Thesis, South

Dakota State University, Brookings.

464

Stonecat, Noturus flavus (Rafinesque, 1818)

Etymology and Synonyms: Noturus = “back tail”, referring to the fusion of the adipose fin with the caudal fin; flavus = “yellow”.

Description: Body elongate; cylindrical anteriorly, laterally compressed posteriorly.

Dorsally tan to dark brown with a cream-white spot at the rear of the dorsal fin base; laterally brown to tan with no distinct markings; ventrally cream to yellow; ventral fins with a dark base, fading to a pale yellow on distal ends; caudal fin often with horizontal stripe at base. Head broad, slightly dorsoventrally compressed. Snout bluntly pointed.

Eyes small, positioned on dorsal side of head. Mouth subterminal, small and wide; upper jaw protrudes past lower jaw. Barbels present; 8 on head surrounding mouth and posterior nostrils. Lips fleshy; upper lip overhangs lower lip. Teeth small, sharp, and numerous on upper and lower jaws; tooth patch present on upper jaw. Gill rakers 6-7.

Dorsal fin with 1 short venomous spine anteriorly followed by 6-7 rays. Adipose fin present; long, short, fused with caudal fin and separated by a shallow notch. Caudal fin square-shaped with nearly straight distal end. Anal fin rays 15-18. Pelvic fins with 8-10 rays; insertions posterior to insertion of dorsal fin. Pectoral fins with single, short venomous spine and 10 rays; spine with serrae on anterior edge, smooth on posterior edge. Lateral line incomplete. Scaleless. Juveniles and spawning adults similar to non- spawning adults.

Similar Species: Closely resembles the Tadpole Madtom. Tadpole Madtom similarly possess an adipose fin fused with the caudal fin, however the caudal fin has a more rounded distal end, unlike the square-shaped caudal fin of the Stonecat. Tadpole Madtom also have jaws that are nearly equal in length, and have eyes placed laterally on the head. 465

Distinguished from other Ictalurid species in the region by the fusion of the adipose fin with the caudal fin.

Distribution and Habitat: Native throughout the north-central United States from

Montana and Wyoming in the northwest, east throughout the Missouri, upper Mississippi,

St. Lawrence-Great Lakes and Ohio River drainages to New York, and south to

Tennessee and Oklahma.1,10,12 Primarily distributed throughout the main tributaries west of the Missouri River in the Dakotas, but is also present within the Red River, James,

Vermillion, and Big Sioux watersheds. Inhabits a wide variety of fluvial cool and warm- water systems, including medium to large size streams and lakes with clear to slightly turbid water and sand, pebble, gravel, or rocky substrates.5,14,15 Often associated with shallow shoreline areas ranging 0.03 m-0.46 m (0.10-4.51 ft) in depth and moderate water velocities ranging 0.00-068 m/s in run and riffle sequences of streams.2,4 Generally avoids areas 0-5 m (0-16.40 ft) away from the nearest shoreline.5 Shoreline areas are likely utilized as they provide opportunities for the species to feed on aquatic invertebrates that drift and become available with increased flows.4 Crevices between coarse substrates are likely used as cover during the day, since they are considered nocturnal like other Ictalurid species.4,6,7 Stonecats are also known to select the shallowest available depths at night.5 Susceptible to dewatering, and avoids intermittent and high gradient streams.5 Similar to adults, juveniles are often associated with coarse substrates, however they tend to avoid shoreline areas, likely because they utilize the midchannel to migrate to shallower habitats.5

Reproduction: Spawning takes place May to July, or when water temperatures reach 17

°C (62.69 °F).4,15 Spawning typically persists until water temperatures hover 25-27 °C 466

(77-80.6 °F).4,15 Information regarding spawning habitat is limited. It is suggested that they inhabit shallow, warm, shorelines areas with large substrates prior to spawning, and shift to deeper waters to spawn.4 Warmer water temperatures near shorelines may increase their metabolism and growth which in turn enhances their reproduction.4 Sexual maturity in females occurs at age 3-4 at an average length of 119.4 mm (4.70 in) SL.15

Males believed to reach sexual maturity at roughly 87 mm (3.42 in).15 Males defend and construct nests by moving small particles with their mouths.8 Nests are generally constructed within pools or between crevices under rocks with aquatic vegetation present at depths 53-117 cm (1.74-3.84 ft) deep.8,15 Fecundity increases with the size of female.15

Fecundity ranges 189-570 eggs per female with approximately 104-306 eggs laid per clutch.15 Eggs approximately 2.6-4.0 mm (0.10-0.16 in) in diameter, demersal, opaque- yellow in color, and held together by an adhesive jelly-like mixture.3,13,15 Males guard and defend the nests until larvae reach approximately 12-15 mm (0.47-0.59 in) TL when they become able to swim individually.6

Age and Growth: Larvae roughly 6.7-7.5 mm (0.26-0.30 in) TL at hatching.15 Growth rates greatest during the first year of life.15 Mean lengths-at-age of individuals from South

Dakota are reported as: age-1, 78.5 mm (3.09 in); age-2, 96.8 mm (3.81 in); age-3, 113.8 mm (4.48 in); age-4, 137.6 mm (5.42 in); age-5, 155.0 mm (6.10 in); age-6, 175.6 mm

(6.91 in); age-7, 193.0 mm (7.60 in) TL.7 Maximum length is often greater and life expectancy is typically longer in lentic environments compared to lotic environments.7,9,12 Capable of reaching 312 mm (12.28 in) TL.12 Longevity up to 9 years.9 467

Food and Feeding: Nocturnal, bottom dwelling, opportunistic predator. Seeks food along the bottom substrate using their barbles to help their sense of smell and taste.11

Consume a diversity of prey items, with some shift relative to body size.1,15 Smaller individuals consume a heavy amount of aquatic insect larvae, including mayfly, stonefly, and caddisflies. Adults consume aquatic insect larvae, and larger amounts of mollusks, minnows, crayfish and aquatic vegetation.1,12,15

Literature Cited:

1. Alberta Sustainable Resource Development. 2004. Status of the stonecat (Noturus

flavus) in Alberta. Alberta Sustainable Resource Development, Fish and Wildlife

Division, and Alberta Conservation Association, Wildlife Status Report No. 52,

Edmonton, AB 22pp.

2. Banks, S.M., and R.J. DiStefano. 2002. Diurnal habitat association of the

madtoms, Noturus albater, N. exilis, N. flavater and N. flavus in Missouri Ozarks

streams. The American Midland Naturalist 148:138-145.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Brewer, S.K. D.M. Papoulias, and C.F. Rabeni. 2006. Spawning habitat

associations and selection by fishes in a flow-regulated prairie river. Transactions

of the American Fisheries Society 135:763-778.

5. Brewer, S.K., and C.F. Rabeni. 2008. Seasonal and diel habitat shifts by juvenile

ictalurids in a flor-regulated prairie river. The American Midland Naturalist

159:42-54.

6. Burr, B.M., and J.N. Stoeckel. 1999. The natural history of madtoms (genus

Noturus), North American’s diminutive catfishes. Pages 51-101 in E.R. Irwin, 468

W.A. Hubert, C. F. Rabeni, J.H. L. Schramm, and T. Coon, editors. Catfish 2000:

proceedings of the International Ictalurid Symposium. American Fisheries

Society, Bethesda, Maryland.

7. Carlson, D.R. 1966. Age and growth of the Stonecat, Noturus flavus Rafinesque,

in the Vermillion River. Proceedings of the South Dakota Academy of Science

45:131-137.

8. Cochran, P.A. 1996. Cavity enhancement by madtoms (genus Noturus). Journal

of Freshwater Ecology 11:521-522.

9. Gilbert, C.R. 1953. Age and growth of the yellow stone catfish, Noturus flavus

(Rafinesque). M.S. Thesis, Ohio State University, Columbus, Ohio.

10. Jenkins, R.E. and N.M. Burkhead. 1993. Freshwater fishes of Virginia. American

Fisheries Society, Bethesda, Maryland.

11. Pflieger, W.L. 1975. The fishes of Missouri. Missouri Department of

Conservation. 343 p.

12. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

13. Simon, T.P., and B.M. Burr. 2004. Description of developmental stages of the

stonecat, Noturus flavus and the slender madtom, Noturus exilis (siluriformes:

ictaluridae). Proceedings of the Indiana Academy of Science 113:123-132.

14. Trautman, M.B. 1981. The fishes of Ohio with illustrated keys, revised edition. Ohio

State University Press, Columbus, Ohio.

15. Walsh, S.J., and B.M. Burr. 1985. Biology of the Stonecat, Noturus flavus

(Siluriformes, Ictaluridae), in central Illinois and Mississippi streams, and 469 comparison with Great Lakes populations and congeners. Ohio Journal of Science

85:85-96.

470

Tadpole Madtom, Noturus gyrinus (Mitchill, 1817)

Etymology and Synonyms: Noturus = back tail, referring to the merging of the adipose fin and caudal fins; gyrinus = Greek for the word tadpole, referring to the similar body shape the species has with a tadpole.

Description: Body stocky and round anteriorly, compressed posteriorly; posterior end angled downward giving a curved or bent appearance. Dorsal and lateral sides of head, body, and caudal peduncle dark brown to olive grey; ventral side of head and body light yellow-brown to tan with dark, narrow mid lateral stripe extending to caudal fin base; skin thin, sometimes transparent; scales completely absent; fins typically dark brown with paired fins light to medium tan in color; no patterns. Head wide; somewhat depressed dorsally. Mouth wide, terminal with fleshy lips; upper and lower jaw about equal in length; 4 dark colored barbels present, 2 outer and 2 inner positioned in a transverse line on the ventral side of head; teeth small and sharp, positioned on inside margin of upper and lower jaws. Eye small, positioned lower on head. Dorsal fin origin slightly anterior to the midpoint between pectoral and pelvic fins; anterior base swollen with single, short spine followed typically by 6-7 rays; dorsal adipose fin present, short and fused with caudal fin. Caudal fin homocercal and broadly rounded; slightly pointed in in young.

Anal fin somewhat rounded with 13-18 rays; little space between anal fin insertion and caudal fin base. Pelvic fin with 8-10 rays. Pectoral fin with 1 spine, not serrated, roughly two-thirds fin length; with 7-9 rays. Lateral line complete. Poison glands present at base of dorsal and pectoral spines.

Similar Species: The Stonecat (Noturus flavus) also possesses an adipose fin connected to the caudal fin, however the caudal fin is more square in shape, unlike the broadly 471 rounded caudal fin of the Tadpole Madtom. The Stonecat’s upper jaw extends beyond the lower jaw; body shape much more slender anteriorly; eyes are placed higher on the head than the Tadpole Madtom. The Stonecat also displays a horizontal stripe in the center of the caudal fin.

Distribution and Habitat: Endemic to eastern North America; spans from southern

Canada to the Great Lakes and continues along the Mississippi River lowlands to the

Atlantic and Gulf coastal plains.1,7 Known to be absent from the Appalachian Highlands.2

In North Dakota, it has been recorded throughout the James River and all tributaries of the Red River.3,4 Within South Dakota, it has been taken sparingly from the Minnesota,

Big Sioux, and James River drainages east of the Missouri River.5,6 The species prefers habitats of slow-moving, low gradient small streams, sloughs, ponds, and lakes with a silt or mud substrate and heavy amounts of aquatic vegetation and organic debris. They are found within moderately clear to turbid water often within vegetation or along undercut banks during the day to hide from predators. They also have been known to inhabit empty bottles or aluminum cans.10

Reproduction: Little information is known regarding reproductive behaviors of the

Tadpole Madtom due to the species nocturnal and cautious habits. Sexual maturity is said to occur between ages one and two.7,12 Spawning typically takes place during the summer months, sometimes extending into early September.7,8,11 Nests are usually made within natural crevices like empty crayfish holes or under logs and rocks.8,9 They also have been known to create nests in empty bottles or aluminum cans.10 Females are known to produce 48-323 mature ova, with clutch size ranging 47-150 eggs per nest.7 Larger females have been noted to produce a greater amount of mature ova.7 Tadpole Madtom 472 females spawn more than once during the breeding season, unlike female bullheads and catfish who deposit all their eggs at once.13 Eggs are demersal, spherical, adhesive, and range in diameter from 2.8-3.5mm (0.110-0.138in).7 Nest is guarded by one or both parents until eggs hatch. Further parental care after hatching is unknown.

Age and Growth: Adults often do not exceed 75-101.6mm (3-4in) TL, but are capable of reaching 129.54mm (5.1in) TL. Young of year in Minnesota measured 15-35mm (0.59-

1.38in) TL; age 1, 43-85mm (1.69-3.35in) TL; age 2, 78-104mm (3.07-4.09in) TL.14,15

Typical lifespan is 2-3 years, with few individuals surviving to age 4.

Food and Feeding: Feeds primarily at night along the substrate or within aquatic vegetation. Organic debris, chironomids, and crustaceans such as amphipods, cladocerans, and ostracods are heavily preyed upon. They also been known to consume small fishes, dragonflies and worms. Young and smaller in size individuals rely heavily on chironomids, oligiochates, and small fly larvae.

Literature Cited:

1. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

2. Gilbert, C.R., and J.D. Williams. 2002. National Audubon Society Field Guide to

Fishes, Revised Edition, North America. Alfred A. Knopf, Inc., New York.

3. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

4. Russel, G.W. 1975. Distribution of fishes in North Dakota drainages affected by

the Garrison Diversion Project. M.S. Thesis, University of North Dakota. 100p. 473

5. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

6. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

7. Whiteside, L.A., and B.M. Burr. 1986. Aspects of the life history of the Tadpole

Madtom, Noturus gyrinus (Siluriformes: Ictaluridae), in Southern Illinois. The

Ohio Journal of Science, 86(4): 153-160.

8. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

9. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

10. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

11. Lindquist, D.G., P.W. Shute, and J.R. Shute. 1982. Spawning and nest site

selection by the broadtail and tadpole madtoms: utilization of experimental

spawning cover in Lake Waccamaw, N.C. ASIH Program and Abstracts, 62

Annual Meeting, Northern Illinois University, DeKalb.

12. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

13. Menzel, B., & Raney, E. 1973. Hybrid Madtom Catfish, Noturus gyrinus x

Noturus miurus, from Cayuga Lake, New York. The American Midland

Naturalist, 90(1), 165-176. 474

14. Case, B.E. 1970. An ecological study of the tadpole madtom Noturus gyrinus

(Mitchell), with special reference to movements and population fluctuations. M.S.

Thesis University of Manitoba.

15. Hooper, F.F. 1949. Age analysis of a population of the ameurid fish Schilbeodes

mollis. (Noturus gyrinus) Copeia 1:34-38.

475

Flathead Catfish, Pylodictis olivaris (Rafinesque, 1818)

Etymology and Synonyms: Pylodictis = mud fish, referring to the species habitat preference; olivaris = olive colored, referring to the species coloration.

Description: Body elongate, moderately dorsoventrally compressed anteriorly.

Coloration varies widely with habitat and size. Dorsally and laterally light brown, olive, or light green-yellow with dark black or brown mottling; ventrally yellow, tan, or cream; fins dark, dusky in color, with upper lobe of caudal fin white to cream; barbles light to dark brown-olive; mottling and white edge on upper caudal lobe tend to fade with age.

Head broad, dorsoventrally flattened. Snout broad, blunt. Eye small, positioned dorsally on head. Mouth wide, superior; lower jaw extends past upper jaw. Barbels present, four pairs total; one pair on top of snout; one long pair near corners of mouth on upper jaw; two pairs on chin, with inner pair shorter outer pair. Teeth present on upper and lower jaws, small and sharp; premaxillary tooth patch present on upper jaw with backwards extensions. Gill rakers 9-13. Dorsal fin with 1 short spine, 6 rays. Adipose fin present, elongate, flap-like. Caudal peduncle slender, short. Caudal fin slightly forked, almost square in shape. Anal fin directly under adipose fin; rounded distal end with 14-17 rays.

Pelvic fin small, abdominal with 8-9 rays. Pectoral fin with 1 strongly serrated spine on anterior and posterior edges; >1/2 the fin length. Lateral line complete. Scaleless.

Juveniles similar to adults with prominent white or cream upper caudal lobe. Spawning adults similar to non-breeding adults.

Similar Species: Juveniles and young adults similar in appearance to bullheads, the

Stonecat, or Tadpole Madtom. Bullheads display an anal fin with >17 rays. Stonecat lack distinct mottling and the adipose fin is strongly fused with the caudal fin. Tadpole 476

Madtom also have a fused adipose and caudal fin, as well as a broadly rounded caudal fin. Channel Catfish and Blue Catfish have deeply forked caudal fins and upper jaws extending past the lower jaw.

Distribution and Habitat: Native to the southern Great Lakes region, as well as the

Mississippi, Mobile, and Rio Grande river drainages. Has been widely introduced and stocked outside its native range. Occurs in the Missouri River and its major tributaries in

North and South Dakota. Primarily a large river species, but also inhabits creeks, lakes, and reservoirs. Prefers slow water velocities in dark areas with abundant submerged cover such as rocky coverage, riprap, or woody debris in areas with low gradient.8

Overwinter in deeper main channel pools with riprap and reduced flow rates.4 In

Michigan, Flathead Catfish migrate a mean distance of 1,146 m (3,759 ft) to winter habitat when water reached 10°C (50°F), and migrated a mean distance of 1,045 m (3428 ft) back to their summer habitats in spring when water temperatures reached 10°C

(50°F).4 Primarily nocturnal and exhibit low activity during the day. Young inhabit shallow areas near gravel riffles.2

Reproduction: Spawning takes place June-July when water temperatures reach 22.2-

23.9°C (72-75°F).2,9 Sexual maturity reached at age 3-4. Both sexes prepare a nest, often near stream banks or cavities within submerged structures, by creating depressions in the substrate with their mouths and tails.2 Eggs golden-yellow in color, adhesive. Fecundity increases with size of female and averages 3,520-3,783 eggs/ kg of body weight.3,10

Following fertilization of eggs, the male chases the female out of the nesting area, which he will continue to fan and defend until the larvae hatch.2 Hatching occurs within 6-9 days.2,9 477

Age and Growth: Second largest ictalurid in North America. Larvae roughly 4-11 mm post hatching. Growth in lakes known to be faster than native riverine populations.1,5,8

Growth rates are highest in shallow, turbid, lower portions of streams.9 Mean lengths-at- age from the James River were reported as: age-1, 83 mm (3.27 in) TL; age-2, 186 mm

(7.32 in) TL; age-3, 294 mm (11.57 in) TL; age-4, 385 mm (15.16 in) TL; age-5, 468 mm

(18.43 in) TL; age-6, 539 mm (21.22 in) TL; age-7, 599 mm (23.6 in) TL; age-8, 650 mm

(25.6 in) TL; age-9, 707 mm (27.8 in) TL; age-10, 755 mm (29.7 in) TL; age-11, 793 mm

(31.2 in) TL.1 Capable of exceeding 1000 mm (39 in) TL, and 50 kg (110 lbs). Longevity

28 years, although fish over 13 are rare in most populations.7

Food and Feeding: Majority of feeding takes place at night, when individuals migrate from deep to shallow areas near riffles. Adults primarily piscivorous, apex, opportunistic predators. Popular forage fish include Gizzard Shad, Freshwater Drum, White Perch and

Common Carp. Juveniles primarily invertivores. In Lake Mitchell, South Dakota, individuals <300 mm (11.8 in) TL mainly consumed crayfish; individuals 300-600 mm

(11.8-23.6 in) TL consumed nearly equal amounts of crayfish and smaller prey fish

(centrarchids), and large adults >600 mm (23.6 in) TL consumed only prey fish.8 A diet shift to piscivory occurs at approximately 400 mm (15.7 in) in Lake Mitchell, and at 200-

350 mm (7.8-13.7 in) TL in the Missouri River, but size-at-shift may depend on availability and production of prey.6,8,9

Literature Cited:

1. Arterburn, J.E. 2001. Population characteristics and sampling methods of catfish

for the James and Big Sioux rivers. M.S. Thesis. South Dakota State University,

Brookings. 478

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Colehour, J.D. 2009. Fecundity of flathead catfish and blue catfish from the

Mississippi River between Hannibal and Cape Girardeau, Missouri. M.S. Thesis,

University of Central Missouri, Warrensburg.

4. Daugherty, D.J., and T.M. Sutton. 2005. Seasonal movement patterns, habitat use,

and home range of Flathead Catfish in the Lower St. Joseph River, Michigan.

North American Journal of Fisheries Management 25:256-269.

5. Goble, C.W. 2011. Ecology and management of channel catfish Ictalurus

punctatus and flathead catfish Pylodictis olivaris populations in the Missouri

River, NE. M.S. Thesis, University of Nebraska, Lincoln.

6. Hogberg, N.P., and M.A. Pegg. 2016. Flathead catfish Pylodictis olivaris diet

composition during extreme flow events in a large river. Journal of Freshwater

Ecology 31(3)431-441.

7. Kwak, T.J., W.E. Pine III, and D.S. Waters. 2006. Age, growth, and mortality of

introduced flathead catfish and a review of other populations. North American

Journal of Fisheries Management 26:73-87.

8. Lucchesi, D.O., M.D. Wagner, T.M. Stevens, and B.D.S. Graeb. 2017. Population

dynamics of introduced flathead catfish in Lake Mitchell, South Dakota. Journal

of Freshwater Ecology 32(1):323-336.

9. Minckley, W.L., and J.E. Deacon. 1959. Biology of flathead catfish in Kansas.

Transactions of the American Fisheries Society 88:344-355.

10. Summerville, V.C., and H.D. Crawley. 1970. Egg production of flathead catfish.

The Progressive Fish-Culturist 32(4):191-191. 479

CHAPTER 13

FAMILY ESOCIDAE

Introduction

The Pike family, represented by the genus Esox, contains a total of five species worldwide, with four species native to North America. The Amur Pike, Esox reichertii, does not occur within the United States, as it is endemic to China, Mongolia and Russia within the Amur River basin. The Northern Pike, Esox lucius, has a circumpolar distribution in that is native to North America, but is also native to Asia and Europe.

Three species within the genus Esox occur within the Dakotas: the Northern Pike, the

Muskellunge, Esox masquinongy, and the Grass Pickerel, Esox americanus. Elongate, cylindrical, torpedo-like bodies represent members of the genus Esox, as well as an elongate, dorsoventrally flattened, duckbill-like snout. Esox species are also recognized by having large, sharp canine teeth, a complete lateral line, no adipose fin, and dorsal and anal fins that are set far posterior on the body. Although Esox species are commonly recognized by their coloration and markings, these features sometimes vary making it hard to tell one species apart from another. Within the genus Esox, species can more easily be distinguished from others within the genus by the scaling on the opercle, and the mandibular pores located on the underside of the snout. For example, the opercle of a

Muskellunge and Northern Pike is only scaled on the top half, whereas it is fully scaled on the Grass Pickerel. Northern Pike have 5 mandibular pores present on each side of the jaw, whereas Muskellunge have 6-9, and Grass Pickerel have 4. 480

Pikes occur within lakes, reservoirs, sloughs, streams and rivers, and prefer areas with little to no flow. They are primarily piscivorous, although they are capable of feeding opportunistically on other prey items in response to changes within their environment. Pikes are also visual and diurnal feeders that often exemplify ambushing behavior on their prey. Spawning takes place in the spring, and a single female often spawns with one or more males. Pikes do not construct any sort of nest for the eggs, and no parental care is given pre- or post- spawning. The Northern Pike and Muskellunge are capable of reaching large sizes and are well sought after game species within the

Dakotas.

The family Esocidae also includes the Mudminnows, which belonged to the past family Umbridae. Now merged into the same family as Pikes, the Mudminnows are represented by the genera Umbra, which includes the Central Mudminnow, Umbra limi, found in the Dakotas.

481

Grass Pickerel, Esox americanus vermiculatus (Lesueur, 1846)

Etymology and Synonyms: Esox = an old European vernacular name for the pike; americanus = Latin for “of America”; vermiculatus = “vermiculated”, referring to the wavy, oblique vertical bars or vermiculations on the lateral sides. The Grass Pickerel is a subspecies of the Redfin Pickerel, Esox americanus, which is the same morphologically, but occurs separately geographically.8

Description: Body elongate and cylindrical. Dorsally dark green to olive; laterally olive green with darker green patches forming wavy, oblique vertical bars; ventrally cream to white; fins generally clear with no markings but may have slight gold to cream coloration present; dark brown to black bar present below eye, slanting slightly posterior. Head elongate. Opercle fully scaled; cheek fully scaled. Snout elongate, flattened dorso- ventrally. Eye relatively large, placed laterally on the upper portion of the head. Mouth terminal and large; upper jaw does not extend past the middle of the eye. Teeth present on lower jaws; large, and canine-like. Mandibular pores present; 4 or less on each ventral side of the lower jaw. Branchiostegal rays usually 12 or less on each side of jaw. Gill rakers sharp. Dorsal fin set far posterior on body with nearly rounded distal end; 12-13 rays. Adipose fin absent. Caudal peduncle elongate and thick. Caudal fin moderately forked with slightly rounded lobes. Anal fin with 11-12 rays; insertion directly inferior to insertion of dorsal fin. Pelvic fins abdominal with 9-11 rays. Pectoral fins with 14-15 rays and rounded distal ends. Lateral line complete with 97-110 cycloid scales in series.

Juveniles have a rather distinct light, pale lateral band that breaks up and blends in as the fish grows. 482

Similar Species: Although Grass Pickerel attain much smaller maximum lengths, they closely resemble the Northern Pike and the Muskellunge. Grass Pickerel can easily be distinguished from these two species by the presence of the brown to black, dark, vertical bar located underneath the eye. Furthermore, Northern Pike have an opercle that is only scaled on the top half, roughly 8 horizontal rows of small, light yellow to cream colored bean-shaped spots present on the lateral sides, teeth present on both jaws, and 5-6 mandibular pores present on each ventral side of the lower jaw. Muskellunge also have an opercle that is only scaled on the top half, have 6-9 mandibular pores present on each ventral side of the lower jaw, and have faint, dark vertical bars or irregular spots on the lateral sides.

Distribution and Habitat: Grass Pickerel are native west of the Appalachian Mountains within the Mississippi and Great Lakes draianges.5 Grass Pickerel are not known to occur in North Dakota. In South Dakota, the species has only been documented within the

Missouri River basin below Lake Francis Case in the very southeastern corner of the state. Grass Pickerel inhabit clear to slightly turbid shallower waters, generally <2 m

(6.56 ft) in depth, in areas with little to no flow in sloughs, ponds, small streams, lakes and shallower, sluggish bays of larger bodies of water with silt and mud substrates.1,3,7

The species is very reliant on abundant emerged, floating, or submerged aquatic vegetation.3

Reproduction: Spawning occurs during the spring and is generally initiated by increasing water levels and new vegetative growth.3 Specific information on the preferred spawning habitat of Grass Pickerel is limited, although it is known that they spawn in sloughs, near shorelines, and within bays near various forms of vegetation.3,7 Sexual 483 maturity may be reached within the first year of life for both males and females.7 Grass

Pickerel are broadcast spawners, so no nest is prepared and no parental care is given.3

Eggs are demersal, adhesive to vegetation, clear to yellowish-amber in color, and roughly

1.5-2.4 mm (0.06-0.09 in) in diameter.1,3 Eggs hatch within 11-15 days in water temperatures 7.8-8.9 °C (46.04-48.02 °F).1

Age and Growth: Grass Pickerel are the smallest members of the genus Esox in the

Dakotas. Mean lengths-at-age of Grass Pickerel from Pony Lake in the Nebraska

Sandhills region are reported as: age-2, 166 mm (6.54 in) TL; age-3, 172 mm (6.77 in)

TL; age-4, 178 mm (7.01 in) TL; age-5, 224 mm (8.82 in) TL.6 Growth rates from Pony

Lake are known to be substantially slower than rates from other populations, likely due to the a higher abundance of Common Carp and a reduced amount of submerged aquatic vegetation.2,6,7 Mean lengths-at-age from multiple lakes in Wisconsin are reported as: age-0, 145 mm (5.71 in) TL; age-1, 208 mm (8.19 in) TL; age-2, 251 mm (9.88 in) TL; age-3, 287 mm (11.30 in) TL; age-4, 356 mm (14.02 in) TL.1 Female Grass Pickerel generally exceed males in length at each age and longevity.7 Capable of reaching 381 mm

(15.00 in) TL.10 Longevity 5 years.1,9

Food and Feeding: Like other Esocid species, Grass Pickerel are ambush predators and feed by sight primarily in the afternoon and evening.1 Grass Pickerel display opportunistic foraging behavior, and are known to undergo and ontogenetic diet shift with increasing size, as well as exemplify a focus on consuming larger prey items with increasing Grass Pickerel length.7,11 In Indiana, fishes were the dominant prey item of

Grass Pickerel 57-95 mm (2.24-3.74 in) TL, fishes and crayfish were the dominant prey items of Grass Pickerel 96-150 mm (3.78-5.91 in) TL, and crayfish were the dominant 484 prey item of individuals >150 mm (5.91 in) TL.11 In Wisconsin, Grass Pickerel 9.5-15 mm (0.37-0.59 in) TL primarily consumed zooplankton such as Cladocerans, Copepods and Ostracods, while diets of individuals 15-40 mm (0.59-1.57 in) TL included

Copepods, Cldocerans, Tendipedid larvae, Odonata nymphs and fish.7 Grass Pickerel are capable of consuming large food fish for their size; for instance, a Wisconsin study reported a 244 mm (9.61 in) TL Grass Pickerel containing a 102 mm (4.02 in) Bluegill in its stomach, as well as a 198 mm (7.80 in) TL Grass Pickerel with a 76 mm (2.99 in) TL

Yellow Perch in its stomach.7 Adult Grass Pickerel are known to be a cannibalistic, and smaller Grass Pickerel have been found in the stomachs of Grass Pickerel fingerlings.3,7,4

Other prey items ingested by Grass Pickerel include Coleopternas, Ephemeropterans,

Annelids and occasionally frogs.11

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Carlander, K.D. 1969. Handbook of freshwater fishery biology, volume 1. Iowa

State University Press, Ames.

3. COSEWIC. 2005. COSEWIC assessment and status report on the grass pickerel

Esox americanus vermiculatus in Canada. Committee on the Status of

Endangered Wildlife in Canada. Ottawa. 27 pp.

4. Crossman, E.J. 1962. The grass pickerel Esox americanus vermiculatus LeSueur

in Canada. Royal Ontario Museum, Life Sciences Division. 55:1-29.

5. Crossman, E.J. 1966. A taxonomic study of Esox americanus and its subspecies in

eastern North America. Copeia, 1966: 1-20. 485

6. Jolley, J.C., and D.W. Willis. 2008. Characteristics of a Grass Pickerel (Esox

americanus vermiculatus) population in Pony Lake, Nebraska. Journal of

Freshwater Ecology 23:497-499.

7. Kleinert, S.J., and D. Mraz. 1966. Life history of the grass pickerel (Esox

americanus vermiculatus) in southeastern Wisconsin. Wisconsin Conservation

Department Technical Bulletin 37, Madison.

8. Reist, J.D., and E.J. Crossman. 1987. Genetic basis of variation in morphometric

characters as implied by hybrids between subspecies of Esox americanus (Pisces:

Esocidae). Canadian Journal of Zoology 63:1429-1439.

9. Serns, S.L., and T.C. McKight. 1977. The occurrence of Northern Pike x Grass

Pickerel hybrids and an exceptionally large Grass Pickerel in a northern

Wisconsin stream. Copeia 1977:780-781.

10. Trautman, M.B. 1957. The fishes of Ohio. Ohio State University Press,

Columbus. 683pp.

11. Weinman, M.L., and T.E. Lauer. 2007. Diet of Grass Pickerel (Esox americanus

vermiculafus) in Indiana streams. Journal of Freshwater Ecology 22:451-460.

486

Northern Pike, Esox lucius (Linnaeus, 1758)

Etymology and Synonyms: Esox = an old European vernacular name for the pike; lucius

= Latin name for pike.

Description: Body elongate, cylindrical. Dorsally dark green to olive; laterally olive with roughly 8 horizontal rows of small, bean-shaped light yellow to cream colored spots; ventrally cream to white; fins gold to amber in color with dark spots or blotches present.

Head elongate. Opercle only scaled on top half; cheek fully scaled. Snout elongate, flattened dorso-ventrally. Eye positioned high on head. Mouth terminal and large. Teeth present on both jaws, roof of mouth, tongue, and branchial bones; lower jaw with large, sharp teeth, and tongue and roof of mouth with short, comb-like teeth. Mandibular pores present; 5 on each ventral side of lower jaw. Branchiostegal rays 14-16 on each jaw. Gill rakers sharp. Dorsal fin set far posterior on body with round distal end; 15-19 rays.

Adipose fin absent. Caudal peduncle elongate, thick. Caudal fin moderately forked with rounded lobes. Anal fin with 12-15 rays. Pelvic fin abdominal with 10-11 rays. Pectoral fins with 14-17 rays and rounded distal end. Lateral line complete with 105-148 cycloid scales. Juveniles faintly mottled with alternating dark and light vertical bars laterally.

Similar Species: Closely resembles the Grass Pickerel and Muskellunge. Grass Pickerel have a fully scaled opercle, 4 mandibular pores on each side of the jaw, 11-14 branchiostegal rays, a dark vertical bar below the eye, and wavy, vertical dark bars on lateral sides with a thin, light green to cream lateral stripe. Muskellunge with opercle and cheek scaled on the top half only, 6-9 mandibular pores on each side of the jaw, 16-19 branchiostegal rays, and faint, dark vertical bars or irregular spots on lateral sides. 487

Distribution and Habitat: Only species in the esocid family native to North America, northwestern Europe, and northern Asia.11 Native in North America throughout Alaska, most of southern Canada, the Missouri and Mississippi River drainages north of the confluence, the Ohio River drainage east to Vermont, the Great Lakes basin, and south to central Missouri.4 Widely distributed outside of the native range due to popularity as a sportfish. Mesothermal, coolwater species that inhabits a wide variety of habitats including small to medium sized rivers, streams, lakes, and reservoirs. Most frequent in lakes, pools, and backwaters with clear water, little current, and emerged or submerged aquatic vegetation coverage.8 In eastern South Dakota lakes, Northern Pike tolerate summer water temperatures exceeding 25°C (77°F), well above the optimum water temperature of 8-18°C (46.4-64.4°F), indicating the adaptability of the species.6,7 Adults occupy shallow water in spring and fall, transitioning to deeper depths in summer and winter, however they are rarely found below the thermocline.11,4 Tolerant to a wide range of chemical and physical conditions. Juveniles heavily dependent on areas with dense aquatic vegetation, such as small wadeable streams, which they utilize as a nursery in the beginning stages of life.5

Reproduction: Spawning takes place in spring, following ice-out, or when water temperatures reach 8-12°C (46.4-53.6°F).4 Adults migrate from deeper water into shallow areas of lakes, or calm backwaters of rivers, over submerged vegetation; without submerged vegetation, reproductive success of the species is low.3 Males enter spawning grounds prior to females. One to two males escort and female over the spawning grounds where eggs are released and fertilized. Males reach sexual maturity ages 1-2; females ages 2-3. No nest is prepared and no parental care is given. Adults return to deeper waters 488 after spawning. Fecundity dependent of size of female, but may exceed 100,000. Eggs amber in color, 2.5-3.0 mm (0.09-0.12 in) in diameter, demersal, and adhesive to submerged vegetation or substrate. Hatching occurs within 14 days.

Age and Growth: Growth is highly variable among and within populations.4 Optimum growth for Northern Pike in South Dakota occurs in water temperatures 8-18°C (46.4-

64.4°F).6,7 Growth is rapid during the first two years. Females often larger and grow faster than males. Large individuals often 609-914 mm (24-36 in) TL.2 Average longevity

8 years with a natural maximum of 30.

Food and Feeding: Visual, opportunistic, ambush predators. Adults primarily piscivorous, but feed opportunistically on a variety of prey items (e.g., leopard frogs).1,5,9

Most active feeding takes place during the day. In Lake Thompson, South Dakota, adults primarily consumed Common Carp, Black and White Crappie, and Fathead Minnows.9

Fathead minnows were an important prey source throughout the year, while consumption of Common Carp and crappies was more seasonal.9 Age-0 individuals from small streams are known to feed on a wide range of vertebrate and invertebrate prey items such as

Diptera, Ephemeroptera, larval common carp, and amhipods.5 Smaller individuals from

Lake Pactola, South Dakota consumed Rainbow Smelt, Rock Bass, Bluegill, and other

Centrarchids.10

Literature Cited:

1. Chapman, L.J., W.C. Mackay, and C.W. Wilkinson. 1989. Feeding flexibility in

Northern Pike (Esox lucius): fish versus invertebrate prey. Canadian Journal of

Fisheries and Aquatic Sciences 46:666-669. 489

2. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

3. Hassler, T.J. 1970. Environmental influences in early development and year-class

strength of Northern Pike in Lakes Oahe and Sharpe, South Dakota. Transactions

of the American Fisheries Society 99(2):369-375.

4. Inskip, P.D. 1982. Habitat suitability index models: Northern Pike. U.S.

Department of Interior, Fish and Wildlife Service FWS/OBS-82/10.17.

5. Krause, J.R. 2013. Biotic integrity and Northern Pike ecology in Eastern South

Dakota. M.S. Thesis, South Dakota State University, Brookings.

6. Neumann, R.M. 1994. Growth, distribution, and movement of Northern Pike in a

South Dakota natural lake, with sampling considerations. Dissertation, South

Dakota State University, Brookings.

7. Neumann, R.M., D.W. Willis, and S.M. Sammons. 1994. Seasonal growth of

Northern Pike (Esox lucius) in a South Dakota glacial lake. Journal of Freshwater

Ecology 9:191-196.

8. Paukert, C.P., and D.W. Willis. 2003. Population Characteristics and ecological

role of Northern Pike in shallow natural lakes in Nebraska. North American

Journal of Fisheries Management 23:313-322.

9. Sammons, S.M. 1993. Annual food habits of Northern Pike in an eastern South

Dakota natural lake. M.S. Thesis, South Dakota State University, Brookings.

10. Scheibel, N.C. 2015. Age, growth, and trophic interactions of Lake Trout and

Northern Pike in Pactola Reservoir: Implications for lake trout management. M.S.

Thesis, South Dakota State University, Brookings. 490

11. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

491

Muskellunge, Esox masquinongy (Mitchill, 1824)

Etymology and Synonyms: Esox = an old European vernacular name for the pike; masquinongy = name applied to the species by the Ojibwae Indians.

Description: Body elongate, cylindrical. Dorsally dark olive; laterally olive to silver with dark, diffuse vertical bars or spots, less apparent in larger adults; ventrally cream to white; fins yellow to amber; dorsal, anal, and caudal fin with large, dark, diffuse spots present. Head elongate. Opercle and cheek only scaled on top half. Snout elongate, flattened dorso-ventrally. Eye positioned high on head. Mouth terminal and large; maxillary bone extending to mid or posterior edge of pupil. Teeth present on both jaws, roof of mouth, tongue, and branchial bones; lower jaw with large, sharp canines; tongue and roof of mouth with short, comb-like teeth. Mandibular pores present; 6-9 on each ventral side of lower jaw. Branchiostegal rays 16-19 on each jaw. Gill rakers sharp, tooth-like. Dorsal fin set far posterior on body with slightly straight distal end; 15-19 rays. Adipose fin absent. Caudal peduncle elongate, thick. Caudal fin moderately forked with slightly pointed lobes. Anal fin with 14-16 rays. Pelvic fin abdominal with 11-12 rays. Pectoral fins with 14-19 rays and slightly straight distal end. Lateral line complete with 147-155 cycloid scales in series. Juveniles with broken dark vertical bars appearing as spots in irregular columns.

Similar Species: Closely resembles the Northern Pike. Northern Pike have an opercle that is scaled on the top half only, but have a fully scaled cheek. Northern Pike also have

5 mandibular pores on each side of the jaw, 14-16 branchiostegal rays, and roughly 8 horizontal rows of small, bean-shaped light yellow to cream colored spots on the lateral sides. 492

Distribution and Habitat: Endemic to North America. Native to the Upper Mississippi,

Great Lakes, Ohio, and Hudson Bay river drainages. Due to poor reproductive success and destruction of spawning habitat, the species is declining throughout its native range.2,3,5 Many populations are maintained by stocking, including populations in eastern

North Dakota and South Dakota. Inhabits lakes, rivers, streams, impoundments, and ponds. Typically occurs at depths less than 5 m (15 ft.), but are sometimes found between

12-15 m (40-50 ft.) in lakes.1 Found within low gradient pools with little current in rivers.

Often associated with submerged vegetation, rock reefs, or structures such as trees.

Optimal water temperature is 25-26°C (77-79°F).6 Establish home ranges, especially during the summer, which may range from 20-500 ha (49-1,235 ac) in size.4 Daily distance traveled is known to increase with an increase in temperature.7 Capable of withstanding low levels of oxygen.1 Fingerlings depend on submerged aquatic vegetation to escape predators.

Reproduction: Spawning begins in spring, typically late April or early May, when water temperatures reach 9.4-15.5°C (49-60°F).6 Spawning habitat depends on location, but generally takes place in water <1 m-3 m (3.3-9.8 ft.) over silt or mud substrates with abundant amounts of debris and dead aquatic vegetation, with gravel, sand or silt substrates.3,6,8 Areas with high dissolved oxygen are required for embryos and larvae to successfully develop.5 Sexual maturity reached at age 3-4 in males; females age 4-5.

Males enter spawning grounds prior to females. A single female is escorted over the spawning ground by one or two males, where their bodies vibrate together to release and fertilize the eggs.2,6 Fractional spawner, with multiple irregular spawning intervals lasting several days.2,6 Eggs amber in color, 2.5-3.5 mm (0.10-0.13 in) in diameter, non- 493 adhesive, and broadcast over vegetated substrate and bebris.6 No nest is built, and no parental care is given. Fecundity increases with size of female, but ranges from 6,000-

265,000.2 After hatching within 8-14 days in water temperatures 12.2-16.7°C (54-62°F), larvae remain on the bottom substrate for another 10 days before moving to deeper water.1,8

Age and Growth: Growth highly variable among populations, and known to be related to the abundance of prey fishes.2,6 Sexual dimorphic growth; females often larger at any given age, grow faster, and live longer than males.2,6 Known to live longer and experience slower growth rates in the northern part of its range.2,6 Growth is most rapid during the first 3 years of life.1 Average 762-1,168 mm (30-46 in) TL.6 Capable of reaching 1,829 mm (72 in) TL, and 31,751.5 g (70 lbs). Long-lived species. Average longevity 10-15 years; maximum 30 years.

Food and Feeding: Large, apex, visual, ambush predator. Adults mainly feed during the day and primarily consume a wide variety of other fish species including Yellow Perch, sunfish, minnows, Northern Pike and even smaller Muskellunge. Crayfish, frogs, mice, and small waterfowl have also been reported in diets. Fry mainly consume larger zooplankton such as cladocerans, and switch to a piscivorous diet once roughly 30-40 mm (11.8-15.7 in) TL.2

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Cook, M.F., and R.C. Solomon. 1987. Habitat suitability index models:

Muskellunge. U.S. Fish and Wildlife Service. Biological Report. 82(10.148). 494

3. Crane, D.P., L.M. Miller, J.S. Diana, J.M. Casselman, J.M. Farrell, K.L.

Kapuscinski, and J.K. Nohner. 2015. Muskellunge and Northern Pike ecology and

management: Important issues and research needs. Fisheries 40(6)258-267.

4. Diana, J.S., P. Hanchin, and N. Popoff. 2015. Movement patterns and spawning

sites of Muskellunge Esox masquinongy in the Antrim chain of lakes, Michigan.

Environmental Biology of Fishes 98(3):833-844

5. Dombeck, M.P., B.W. Menzel, and P.N. Hinz. 1984. Muskellunge spawning

habitat and reproductive success. Transactions of the American Fisheries Society

113:205-216.

6. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

7. Younk, J.A. 1982. Distribution, movement, and temperature selection of adult

Walleye and Muskellunge in a power plant cooling reservoir. M.S. thesis, South

Dakota State University, Brookings.

8. Zorn, S.A., T.L. Margenau, J.S. Diana, and C.J. Edwards. 1998. The influence of

spawning habitat on natural reproduction of Muskellunge in Wisconsin.

Transactions of the American Fisheries Society 127:995-1005.

495

Central Mudminnow, Unbra limi (Kirtland, 1840)

Etymology and Synonyms: Umbra = Latin for shadow, shade, or dark, possibly referring to the species preferred habitat; limi = mud.

Description: Body fusiform to cylindrical; moderately elongate. Dorsally dark brown to olive green; laterally tan to black-brown with roughly 14 irregular dark brown vertical bars; ventrally tan to cream; fins uniformly pale brown; prominent dark vertical bar at base of caudal fin. Head scaled dorsally and laterally. Snout blunt, short. Eyes relatively large, placed laterally on head. Mouth terminal; mandible slightly protrudes maxillary; maxillary extends to anterior edge of pupil. Groove on upper lip non-continuous. Teeth small, villiform, present on premaxilla, roof of mouth, and mandible. Gill rakers 13-15.

Dorsal fin rounded; insertion slightly posterior to insertion of pelvic fins; 13-15 rays.

Adipose fin absent. Caudal peduncle thick. Caudal fin homocercal, rounded. Anal fin rays 7-9. Pelvic fin rays 6-7. Pectoral fin rays 14-16. Lateral line absent; scales in lateral line series 34-37. Scales cycloid. Sexually dimorphic; males with elongated anal fin, almost extending to base of caudal fin; females with short anal fin. Spawning males develop an iridescent purple to blue-green coloration on anal fin.

Similar Species: Closely resembles the Banded Killifish. Banded Killifish have a more elongate body shape, display 12-20 dark greenish-brown vertical bars laterally, have a continuous groove on the upper lip, and lack a dark vertical bar at the base of the caudal fin. Banded Killifish also have 10-12 anal fin rays, and a dorsal fin insertion distinctly posterior to the pelvic fins.

Distribution and Habitat: Native to the upper Mississippi River basin from Quebec to

Manitoba, south to northwestern Tennessee, the Great Lakes tributaries, and the 496 headwaters of the Hudson Bay drainage (Red River).5,11 Common in northern glacial regions.4 Isolated populations occur within the Missouri River drainage in east-central

South Dakota, eastern Nebraska, and western Iowa.5,11 Considered hardy, habitat specialists; inhabit shallow, quiet waters (<1.0 m (3.3 ft) deep) with little to no flow such as shorelines of lakes, wetlands, and pools within small streams. Less common in deeper waters with greater flow. Occurs in areas with moderate to dense amounts of submerged aquatic vegetation or detritus over gravel, sand, silt, and mud substrates. Adapted to be tolerant of hypoxic, isolated, and stagnant conditions with high water temperatures by being facultative air breathers; the swim bladder (connected to the pharynx) assists in aerial respirations, and the species has been observed to gulp air from the surface at all sizes.6,9 Also capable of utilizing air bubbles under the ice in winter months.8,9

Reproduction: Spawning migrations begin in early spring (March-April). Migrate upstream or in lateral movements into adjacent flooded areas with abundant vegetation.10

Spawning takes place in water temperatures near 12.8-15.6°C (55-60°F).1 Information regarding spawning activity is limited. Sexually mature at ages 1-2. Eggs 1.2-1.6 mm

(0.05-0.06 in) in diameter, yellow-orange in color, adhesive, and deposited onto beds of vegetation.10 No nest is built, and parental care is said to be given by the female, who will devour any undeveloped eggs.1,12 Fecundity 220-1489 eggs per female; number of eggs increases with size of female.9,10 Hatching occurs within a week. Young migrate from the breeding site into preferred habitat at approximately 30.0 mm (1.2 in) TL, or earlier depending on the season.

Age and Growth: Newly hatched larvae average 5.0 mm (0.2 in) TL.10 Growth known to be greatest within the first year; average 43.0 mm (1.7 in) TL at age-1.1 In Minnesota, 497 lengths at ages 2-4 were reported as: age-2, 62.0-122.0 mm (2.4-4.8 in) TL; age-3, 91.0-

138.0 mm (3.6-5.4 in) TL; age-4, 110.0-143.0 mm (4.3-5.6 in) TL.7 Rarely exceeds lengths of 140 mm (5.5 in).11 Longevity up to 7-9 years.

Food and Feeding: Primarily a benthic carnivore. Vegetative matter has been seen in the diet, however this is thought to be accidental.2,10 Young primarily consume chironomids, ostracods, copepods, and cladocerans.10 Chironomids known to be the most important food source for fish age 0-1.9 Adults less reliant on small crustaceans, and grow to consume a variety of prey items including aquatic insects (ostracoda, amphipoda, ephemeroptera, odonata, diptera), terrestrial insects (oligiochaeta, diptera, aranae, coleoptera), small mollusks, and sometimes small minnows (Brook Stickleback and

Fathead Minnows).7,9 In aquariums, they have been known to lie motionless, then ambush their prey.4 Known to be active feeders during the winter, with the ability to digest rapidly in cold temperatures.3,9

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Cahn, A.R. 1927. An ecological study of southern Wisconsin fishes; the brook

silversides (Labidesthes sicculus) and the cisco (Leucichthys artedi) in their

relations to the region, 11. Illinois biological monographs 11(1).

3. Chilton, G., K.A. Martin, and J.H. Gee. 1984. Winter feeding: an adaptive

strategy broadening the niche of the central mudminnow, Umbra limi.

Environmental Biology of Fishes 10(3):215-219.

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville. 498

5. Fuller, P.L., L.G. Nico, and J.D. Williams. 1999. Nonindigenous Fishes

Introduced into Inland Waters of the United States. American Fisheries Society,

Special Publication 27, Bethesda, MD. 613pp.

6. Gee, J.H. 1980. Respiratory patterns and antipredator responses in the central

mudminnow, Umbra limi, a continuous, facultative, air-breathing fish. Canadian

Journal of Zoology 58:819-827.

7. Jones, J.A. 1973. The ecology of the mudminnow, Umbra limi, in Fish Lake

(Anoka County, Minnesota). PhD Thesis, Iowa State University, Ames, Iowa.

8. Magnuson, J.J., J.W. Keller, A.L. Beckel, and G.W. Gallepp. 1983. Breathing gas

mixtures different from air: an adaptation for survival under the ice of a

facultative air-breathing fish. Science (Washington, D.C.) 220:312-314.

9. Martin-Bergmann, K.A., and J.H. Gee. 1985. The central mudminnnow, Umbra

limi (Kirtland), a habitat specialist and resource generalist. Canadian Journal of

Zoology 63:1753-1764.

10. Peckham, R.S., and C.F. Dineen. 1957. Ecology of the Central Mudminnow,

Umbra limi (Kirtland). The American Midland Naturalist 58(1):222-231.

11. Schilling, E.G., D.B. Halliwell, A.M. Gullo and J.K. Markowsky. 2006. First

records of Umbra limi (Central Mudminnow) in Maine. Northeastern Naturalist

13(2):287-290.

12. Westman, J.R. 1938. Studies on the reproduction and growth of the bluntnose

minnow, Hyborhynchus notatus (Rafinesque). Copeia 1938(2):57-61.

499

CHAPTER 14

FAMILY SALMONIDAE

Introduction

The salmon, trout, and char family, Salmonidae, consists of roughly 170 species within

11 different genra. Salmonids are native across the Northern Hemisphere; however due to their popularity as a source of food and as sportfish, they have now been widely introduced around the world, including waters in North Dakota and South Dakota. Eight species of salmonids regularly occur throughout the Dakotas however, none are native to either state. Salmonids are cold-water fish species, meaning that they thrive in waters roughly 18°C (65°F) or colder. There are no entirely marine species of salmonids, but many species are capable of completing their entire lifecycle in freshwater. Some species of salmonids are anadromous, meaning that they hatch and evolve in freshwater, migrate out to sea where they spend the majority of their lives, and then migrate back into freshwater systems only to spawn.

Distinguishing characteristics of salmonids include the presence of an adipose fin, an axillary process on the pelvic fin, soft-rayed fins, and numerous small, embedded cycloid scales. The Rainbow Smelt (Osmerus mordax) from the family Osmeridae, looks similar to some species of salmonids where they occur sympatricly in the Dakotas, however they are smaller in size and lack an axillary process. Although it has the name

“trout” in the common name, the Trout-perch (Percopsis omiscomaysus) from the family

Percopsidae, differs from salmonid species by having ctenoid scales and soft spines in the dorsal and anal fins. Coloration can vary widely within this family of fishes, ranging from 500 colorfully patterned fish such as Rainbow Trout (Oncorhynchus mykiss) and Brown Trout

(Salmo trutta) to the more solid silver coloration of the Cisco (Coregonus artedi) and

Lake Whitefish (Coregonus clupeaformis).

Most of the natural cold-water habitats capable of withstanding trout populations in the Dakotas occur within the Black Hills of western South Dakota, where trout were introduced in 1886. Numerous streams and tributaries of the Missouri River in the

Dakotas also contain trout species, however they require annual stockings and only provide seasonal fisheries. Construction of the Missouri River reservoirs between the late

1950’s and early 1960’s created additional cold-water habitats in both North and South

Dakota that now provide fisheries for Chinook Salmon (Oncorhynchus tshawytscha),

Cisco, and Lake Whitefish. In general, trout in the Dakotas prefer cool, clear streams and lakes with high dissolved oxygen levels, and sand, gravel, or rocky substrate and submerged vegetation. Chinook Salmon, Lake Whitefish, and Cisco occurring in the

Missouri River reservoirs prefer the cold, clear, well-oxygenated and deeper pelagic waters.

The well-studied spawning behavior of salmonids is rather miraculous, specifically the migration of Pacific Salmon, which travel long and strenuous journeys.

These species are capable of swimming hundreds, if not thousands of miles from the ocean back to the exact stream where they were first hatched. Chinook Salmon in the

Missouri River reservoirs still mimic this behavior even though they never make it out to the oceans during their lifetime. Instead, during the fall months they retreat back into the spawning stations, such as Whitlocks Bay area on Lake Oahe, or along the shorelines 501 where fish hatchery personnel collect them and help with the process of fertilizing their eggs.

502

Cisco, Coregonus artedi (Lesueur, 1818)

Etymology and Synonyms: Coregonus = “angle-eye”, a term created by Artedi, referring to the Greek meanings of pupil and angle; artedi = referring to the “Father of

Ichthyology”, Petrus Artedi, who was also an associate of Linnaeus. Commonly referred to as “Lake Herring”, which is rather misleading due to it not being a member of the herring family “Clupeidae”.

Description: Body elongate, fusiform, laterally compressed, deepest near dorsal fin insertion. Dorsally gray, olive to blue-green; laterally solid silver with faint blue, purple or pink iridescence; ventrally silver to cream; fins creamy white to light grey without markings. Head small, conical. Snout pointed. Eye moderately large, placed laterally on head. Mouth terminal, oblique; upper and lower jaws equal in length extending to middle of eye. Teeth small, present on both jaws. Gill rakers long, comb-like, 36-64. Dorsal fin with 8-11 rays. Adipose fin present, small, flap-like. Caudal peduncle short, moderately thick. Caudal fin strongly forked. Anal fin with 10-13 rays. Pelvic fins abdominal with

10-12 rays, axillary process present. Pectoral fins with 14-17 rays. Lateral line complete with 70-94 cycloid scales in series. Spawning males develop small tubercles on head and body; females with less numerous and developed tubercles. Juveniles similar in appearance to adults.

Similar Species: Resembles the Lake Whitefish. Lake Whitefish exhibit a blunt, extended snout that overhangs an inferior to subterminal mouth. Skipjack Herring with a slightly superior mouth, and adipose fin absent. Goldeye and Mooneye lack an adipose fin. 503

Distribution and Habitat: Native and broadly distributed across the north-central and eastern United States, the Great Lakes, and throughout majority of Canada to the Arctic

Ocean in the north.18 Threats to the species distribution and population declines in various waters include rising water temperatures, alterations in land-use, and invasive species such as the Rainbow Smelt.3,10,15 Occurs only within the Missouri River reservoirs of the Dakotas.13 Most frequent in the pelagic zone of the hypolimnion in well- oxygenated, cold, clear waters of deep lakes and reservoirs during spring and summer months. Form schools during the day, but disperse at night.17 Intolerant of warm water temperature and low dissolved oxygen concentratuons.9 Young are more tolerant of higher water temperatures than adults.8 Upper lethal temperature for young-of-year is

26°C (78.8°F), and roughly 20°C (68°F) for adults.8,9,6 Young-of-year and adult lower lethal temperature is reported as 0°C (32°F).8,9 Larvae inhabit shallow bays near surface during the first month post hatching, and later migrate farther offshore to deeper waters, but continue to remain near the surface.5,16

Reproduction: Spawning migrations consist of movements made to near-shore, shallower areas of lakes, or upstream of large rivers. By moving and spawning into shallow areas near shorelines, they are able to provide important resource subsidies to nearshore egg predators by transporting energy from the pelagic zone to the shoreline region.21 Spawning takes place at night near the surface over a variety of substrate types during fall, and peaks when water temperature reaches 4°C (39.2°F).2,14 Age at sexual maturity varies with population. Most mature at age 1-2, however age 1-5 is also possible.2,11,19,20 Males arrive at spawning grounds 2-4 days prior to females.4,18 As many as 12 males may initially court a single female to the spawning grounds, but 1-2 males 504 are more common later on.2,4 No nest is constructed and no parental care is given.

Broadcast spawners. Fecundity varies with size of female. Females from Lake Superior

269-356 mm (10.59-14.02 in) TL produced 4,314-10,250 eggs each.2,7 Eggs slightly adhesive and average 1.88 mm (0.07 in) in diameter from Lake Superior and Lake

Michigan.2 Eggs incubate overwinter under the ice until hatching takes place in early spring. Hatching occurs within 37-49 days at 10°C (50°F).2,12

Age and Growth: Information on age and growth limited in the Dakotas region.

Optimum temperature range for normal development of Cisco is roughly 2-8°C (35.6-

46.4°F).2,6 Growth rates highest in summer and early fall.11 Adults average 254-381 mm

(10-15 in) TL, but capable of reaching 558.8 mm (22 in) TL in the Missouri River within the Dakotas. Longevity 10-15 years.

Food and Feeding: Primarily zooplanktivores in offshore pelagic zones. Long gill rakers allow the species to strain out large amounts of small zooplankton such as copepods and cladocerans from the water column. Also known to consume chironomids and fish eggs of its own kind.1,2,7 Cisco stomachs are more full during the day than at night, suggesting that individuals feeding at night forage less efficiently than when in a school.17 Stomach fullness is also known to increase with increased school size indicating that schooling activity increases foraging opportunities.17

Literature Cited:

1. Aku, P.M.K., and W.M. Tonn. 1999. Effects of hypolimnetic oxygenation on the

food resources and feeding ecology of Cisco in Amisk Lake, Alberta.

Transactions of the American Fisheries Society 128:17-30.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 505

3. Bryan, S.D. 1995. Bioenergetics of Walleye in Lake Oahe, South Dakota. M.S.

Thesis, South Dakota State University, Brookings.

4. Cahn, A.R. 1927. An ecological study of the southern Wisconsin fishes. The

brook silverside (Labidesthes sicculus) and the cisco (Leucichthys artedi) in their

relations to the region. Illinois Biological Monographs 11:1-151.

5. Clady, M.D. 1976. Distribution and abundance of larval ciscoes, Coregonus

artedii, and burbot, Lota lota, in Oneida Lake. Journal of Great Lakes Research

2:234-247.

6. Colby, P.J., and L.T. Brooke. 1969. Cisco (Coregonus artedii) mortalities in a

southern Michigan lake. Limnology and Oceanography 14:958-960.

7. Dryer, W.R., and J. Beil. 1964. Life history of the lake herring in Lake Superior.

U.S. Fish and Wildlife Service Fisheries Bulletin 63:493-530.

8. Edsall, T.A, and P.J. Colby. 1970. Temperature tolerance of young-of-the-year

cisco, Coregonus artedii. Transactions of the American Fisheries Society 99:526-

531.

9. Frey, D.G. 1955. Distributional ecology of the cisco (Coregonus artedii) in

Indiana. Invest. Indiana Lakes and Streams 4:177-228.

10. Hartman, W.L. 1973. Effects of exploitation, environmental changes, and new

species on the fish habitats and resources of Lake Erie. Great Lakes Fishery

Commission Report 22.

11. Hile, R. 1936. Age and growth of the cisco, Leucichthys artedi (LeSueur), in the

lakes of the northeastern highlands, Wisconsin. Bulletin of the Bureau of

Fisheries 19:210-317. 506

12. Hinrichs, M.A., and H.E. Booke. 1975. Egg development and larval feeding of the

lake herring, Coregonus artedii LeSueur. Mesuem of Natural History, University

of Wisconsin, Stevens Point, Reports on Fauna and Flora of Wisconsin 10:75-86.

13. Hoagstrom, C.W. 2006. Fish community assembly in the Missouri River basin.

Ph.D. Dissertation, South Dakota State University, Brookings.

14. John, K.R. 1956. Onset in spawning activities of the shallow water cisco,

Leucichthys artedi (LeSueur), in Lake Mendota, Wisconsin, relative to water

temperatures. Copeia 1956:116-118.

15. Lawrie, A.H., and J.F. Rahrer. 1973. Lake Superior: case history of the lake and

its fisheries. Great Lakes Fishery Commission Technical Report 19.

16. MacKay, H.H. 1963. Fishes of Ontario. Ontario Department of Lands and Forests,

Toronto.

17. Milne, S.W., B.J. Shuter, and W.G. Sprules. 2005. The schooling and foraging

ecology of lake herring (Coregonus artedi) in Lake Opeongo, Ontario, Canada.

18. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

19. Smith, S.H. 1956. Life history of lake herring of Green Bay, Lake Michigan. U.S.

Fish and Wildlife Service Fisheries Bulletin 57:87-138.

20. Smith, S.H. 1957. Evolution and distribution of the coregonids. Journal of the

Fisheries Research Board of Canada 14:599-604.

21. Stockwell, J.D., D.L Yule, T.R. Hrabik, M.E. Sierszen, and E.J. Isaac. 2014.

Habitat coupling in a large lake system: delivery of an energy subsidy by an 507 offshore planktivore to the nearshore zone of Lake Superior. Freshwater Biology

59:1197-1212.

508

Lake Whitefish, Coregonus clupeaformis (Mitchill, 1818)

Etymology and Synonyms: Coregonus = “angle-eye”, a term created by Artedi, referring to the Greek meanings of pupil and angle; clupeaformis = “herring shaped”, referring to the shape of the body.

Description: Body elongate, fusiform, laterally compressed. Dorsally dark, pale olive green, bronze or silvery gray; laterally solid silver with bluish sheen; ventrally cream to white; dorsal side of head and upper jaw dark gray; lower jaw silvery white; fins light to dark gray without markings or spots. Head small, conical. Snout blunt, overhanging mouth. Eye moderately large, placed laterally on head. Mouth inferior to subterminal, slightly oblique; upper jaw rarely extends past anterior edge of pupil. Teeth small, present on both jaws. Gill rakers rather short, 24-33. Dorsal fin with 10-12 rays. Adipose fin present, small, flap-like. Caudal peduncle slightly elongate, moderately thick. Caudal fin strongly forked. Anal fin 10-12 rays. Pelvic fins abdominal with 10-12 rays, axillary process present. Pectoral fins with 14-17 rays. Lateral line complete with 74-93 cycloid scales in series. Spawning males develop small tubercles on head and body; females with less numerous and developed tubercles. Juveniles similar in appearance to adults.

Similar Species: Closely resembles the Cisco. Cisco have a more pointed snout and terminal mouth, unlike the inferior to subterminal mouth of Lake Whitefish. Skipjack

Herring with a slightly superior mouth, and adipose fin absent. Goldeye and Mooneye lack an adipose fin.

Distribution and Habitat: Native and broadly distributed throughout the majority of

Canada, north to the Arctic Ocean, and south throughout the north-central and eastern

United States including the Great Lakes.14 Non-native to the Dakotas, but have been 509 stocked into Lake Sakakawea and Lake Oahe.18 Distribution and recruitment known to be negatively affected by the competition and predation of Lake Whitefish larvae by

Rainbow Smelt.7,16 Adults school within cold, oligotrophic waters of lakes, reservoirs, and rivers at depths typically 15-50 m (49.21-164.04 ft.).22 Exhibit diel vertical migrations by occupying deeper depths throughout the day and moving up in the water column during the night, especially in the spring and fall.12 Preferred water temperature

10-14°C (50-57.2°F).6,12 Individuals migrate to deeper waters in the fall where they overwinter until spring.12 Larvae inhabit surface waters immediately after hatching.15

Maximum oxygen consumption and critical swimming speed reached at 12°C (53.6°F).3

A water temperature of 10.1°C (50.18°F), recorded as highest larval survival rate, and juveniles prefer water temperatures 15.5-19.5°C (59.9-67.1°F).9,10 Not tolerant of warm water temperatures; upper lethal limit of adults 26.7°C (80.06°F).11

Reproduction: Spawning migrations to shallow spawning grounds begin in late fall.

Spawning takes place at night during late October through early December in water temperatures <6°C (42.8°F), and lasts for roughly a week.1,13,17 Spawning generally occurs at depths 1-4 m (3.28-13.12 ft.) over clean, sandy, rocky, cobble or boulder substrate in inlets of lakes, streams, or windswept shorelines.1,12 Sexual maturity reached at age 3-6.12 Males arrive at spawning grounds prior to females. One to two males court a female towards the surface of the water where eggs and milt are released during a period of splashing and jumping.2 Broadcast spawners. No nest is constructed and no parental care is given. Adults migrate back to deeper waters post spawning. Fecundity increases with size of female and varies significantly among populations.15 Eggs yellow-orange in color, semi-buoyant, and roughly 2.95 mm (0.12 in) in diameter, however egg size is 510 known to be positively correlated with female body size among populations.4,15 Eggs incubate overwinter and hatch in spring. The timespan from fertilization to median hatch date is inversely related to water temperature, and ranges from 41.7 days at 10°C (50°F) to 182 days at 0.5°C (32.9°F).5 Optimum water temperature range for egg incubation is

3.2-8.1°C (37.76-46.58°F).5

Age and Growth: Information on age and growth limited in the Dakotas region. Largest of the coregonid species. Growth varies widely among populations. Larvae roughly 13-14 mm (0.51-0.55 in) TL at hatching.12 Larval growth rate highest at 18.1°C (64.58°F), and is highly correlated with food availability.9,12 Adults typically 254-381 mm (10-15 in)

TL, but are capable of reaching 736.6 mm (29 in) TL.8 Longevity 12 years.12

Food and Feeding: Adults primarily opportunistic benthivores, consuming large amounts of benthic invertebrates including insects, crustaceans, and mollusks.19,20 Also occasionally consume zooplankton, Mysis, oligochaetes, and small fish such as stickleback and alewives.20 Size of benthic prey consumed increases with fish size, however size of pelagic prey is not known to be related to fish size.19 Zooplankton including large bodied copepods, cladocerans, and rotifers are the dominant prey of larvae.19,21

Literature Cited:

1. Anras, M.L.B., P.M. Cooley, R.A. Bodaly, L. Anras, and R.J.P. Fudge. 1999.

Movement and habitat use by lake whitefish during spawning in a boreal lake:

Integrating acoustic telemetry and geographic information systems. Transactions

of the American Fisheries Society 128:939-952.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 511

3. Bernatchez, L., and J.J. Dodson. 1985. The influence of current speed and

temperature on the swimming capacity of lake whitefish (Coregonus

clupeaformis) and cisco (C. artedii). Canadian Journal of Fisheries and Aquatic

Sciences 42:1522-1529.

4. Booke, H.E. 1970. Speciation parameters in Coregonine fishes: Part I. Egg size.

Part II. Karyotype. Pages 61-66 in Biology of coregonid fishes, C.C. Lindsey, and

C.S. Woods, editors. University of Manitoba Press, Winnipeg.

5. Brooke, L.T. 1975. Effect of different constant incubation temperatures on egg

survival and embryonic development in lake whitefish (Coregonus clupeaformis).

Transactions of the American Fisheries Society 104:555-559.

6. Christie, G.C., and H.A. Regier. 1988. Measures of optimal thermal habitat and

their relationship to yields of four commercial fish species. Canadian Journal of

Fisheries and Aquatic Sciences 45:301-314.

7. Crowder, L.B. 1980. Alewife, rainbow smelt and native fishes in Lake Michigan:

competition or predation? Environmental Biology of Fishes 5:225-233.

8. Dryer, W.R. 1963. Age and growth of the whitefish in Lake Superior. Fishery

Bulletin 63:77-95.

9. Edsall, T.A. 1999a. The growth-temperature relation of juvenile lake whitefish.

Transactions of the American Fisheries Society 128:962-964.

10. Edsall, T.A. 1999b. Preferred temperature of juvenile lake whitefish. Journal of

Great Lakes Research 25:583-588. 512

11. Esdall, T.A., and D.V. Rottiers. 1976. The temperature tolerance of young-of-the-

year lake whitefish, Coregonus clupeaformis. Journal of the Fisheries Research

Board of Canada 33:177-180.

12. Gorsky, D. 2011. Lake whitefish (Coregonus clupeaformis) habitat utilization,

early life history, and interactions with rainbow smelt (Osmerus mordax) in

northern Maine. Ph.D. dissertation, University of Maine, Orono.

13. Hart, J.L. 1930. The spawning and early life history of the whitefish, Coregonus

clupeaformis (Mitchill) in the Bay of Quinte, Ontario. Contributions in Canadian

Biology and Fisheries 6:165-214.

14. Hubbs, C., and K. Lagler. 2004. Fishes of the Great Lakes Region. University of

Michigan Press, Ann Arbor.

15. Ihssen, P.E., D.O. Evans, W.J. Christie, J.A. Reckahn, and R.L. DesJardine. 1981.

Life history, morphology, and electrophoretic characteristics of five allopatric

stocks of lake whitefish (Coregonus clupeaformis) in the Great Lakes region.

Canadian Journal of Fisheries and Aquatic Sciences 38:1790-1807.

16. Loftus, D.H., and P.F. Hulsman. 1986. Predation on larval lake whitefish

(Coregonus clupeaformis) and lake herring (C. artedii) by adult rainbow smelt

(Osmerus mordax). Canadian Journal of Fisheries and Aquatic Sciences 43:812-

818.

17. Nester, R.T., and T.P. Poe. 1984. Predation on lake whitefish eggs by longnose

suckers. Journal of Great Lakes Research 10:327-328. 513

18. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

19. Pothoven, S.A., and T.F. Nalepa. 2006. Feeding ecology of lake whitefish in Lake

Huron. Journal of Great Lakes Research 32:489-501.

20. Reckahn, J.A. 1970. Ecology of young lake whitefish (Coregonus clupeaformis)

in South Bay, Manitoulin Island, Lake Huron. Pages 437-460 in Biology of

coregonid fishes, C.C. Lindsey, and C.S. Woods, editors. University of Manitoba

Press, Winnipeg.

21. Teska, J.D., and D.J. Behmer. 1981. Zooplankton preference of larval lake

whitefish. Transactions of the American Fisheries Society 110:459-461.

22. Walden, H. 1964. Familiar freshwater fishes of America. Harper and Row, New

York.

514

Cutthroat Trout, Oncorhynchus clarkii spp. (Richardson, 1836)

Etymology and Synonyms: Oncorhynchus: Greek, onyx, -ychos = nail + Greek, rhyngchos = snout; clarkia: in honor of William Clark, from the Lewis and Clark expedition.

Description: Body slender, elongated, slightly laterally compressed. Dorsally greenish blue to dusky gray; laterally yellow to silver; dark gray to black round spots present on dorsal and lateral sides, noticeably more numerous on posterior end; dorsal fin, adipose fin, and caudal fin heavily spotted; pectoral, pelvic and anal fins brown to red in color with possible spots; reddish-orange slash mark on each underside of lower jaw. Head small to moderate. Snout short, blunt. Eye moderately large, sometimes spotted. Mouth small, terminal; maxillary extending beyond posterior edge of eye. Frenum absent. Teeth small, sharp, present on posterior end of tongue and jaws. Gill rakers 17-21. Dorsal fin with 8-11 rays, no spines. Adipose fin present; small and slender. Caudal peduncle short, thick. Caudal fin slightly forked. Anal fin with 8-12 rays, no spines; distal end rounded.

Pelvic fin with 9-10 rays; small axillary process present. Pectoral fin with 12-15 rays.

Lateral line complete with 130- 240 small cycloid scales in series.

Similar Species: A polytypic species, Cutthroat Trout exist in several geographical specific forms, or subspecies, with a great amount of genetic diversity.5 Cutthroat Trout are easily distinguishable from other trout species in the Dakotas by the red-orange slash marks on each underside of the lower jaw, as well as the many black spots toward the posterior end of the body. Subspecies of Cutthroat Trout are identified by the location and size of the spots along the body. Brook Trout lack red slash marks on underside of jaw, and display red or pink spots outlined in blue on lateral sides. Rainbow Trout have a 515 pinkish-red lateral stripe and have black spots that are more evenly distributed across the body. Rainbow Trout often hybridize with Cutthroat Trout to produce “cutbows”. These hybrids can be distinguished by the basicranchial teeth on the tongue. Brown Trout generally lack dark spots on the caudal fin, and have dark spots outlined in light blue or gray along lateral sides.

Distribution and Habitat: Distributed widely outside its native range due to sport fishing. Native to the Pacific coast from southern Alaska to northern California, and west to central Canada throughout the Rocky Mountain region, south to New Mexico. Native to the headwaters of the Missouri, Platte, Colorado, and Rio Grande river drainages.2

Introduced, rare, and exotic in the Dakotas; found west of the Missouri River in both

North Dakota and South Dakota, however records exist from the Souris and eastern portion of the Missouri River basins in North Dakota. Prefer headwater sections of riverine habitats with clear, cold, swift moving water, over rocky substrates. Often associated with areas rich in cover and well vegetated stream banks. Prefers water temperatures around 14.5-16°C (58-60°F). Able to withstand short periods of time in water temperatures as high as 26°C (78.8°F), although rarely present in waters exceeding

22°C (71.6°F).1,5

Reproduction: Spawning in the early spring, or when water temperatures reach roughly

10°C (50°F) within stream beds over gravel substrate. Length is known to be more influential than age when determining sexual maturity.4 Sexual maturity of males reached at 110-160 mm (4.33-6.23 in) FL; females at 150-180 mm (5.90-7.08 in) FL.4 Females use their caudal fin to create small depressions or “redds” within the gravel to deposit the eggs. During this time, males will fertilize the eggs. Fecundity ranges from 500-1500 516 eggs each.2 Length-fecundity relationships differentiate between populations.4 Eggs hatch within 28-40 days depending on water temperature, and can take as long as 49 days.3,5,6

Fry reside in the rocky, gravel substrate for roughly two weeks after hatching, and then move into deeper, faster water to continue growth.6

Age and Growth: Subspecies and different populations of Cutthroat Trout can average lengths of 304-406 mm (12-16 in). Older individuals may reach 508 mm (20 in). Optimal growth temperature known to be 14.5°C (58°F). Average lifespan is 4-7 years; capable of living up to age-8.

Food and Feeding: Opportunistic feeders. Known to defend feeding territories often in or downstream of riffle habitat. Fry feed on zooplankton. Juveniles and younger adults feed primarily on larval, pupal, and adult forms of aquatic and terrestrial insects such as mayflies, stoneflies, caddisflies and dipterans. Larger fish prey on other smaller fish, fish eggs, terrestrial insects, snails, small rodents, and annelids.

Literature Cited:

1. Behnke, R.J., and M. Zarn. 1976. Biology and management of threatened and

endangered western trout. U.S. Forest Service General Technical Report. RM-28.

45 pp.

2. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

3. Cope, O.B. 1957. The choice of spawning sites by cutthroat trout. Proc. Utah

Academy of Science, Arts, and Letters 34:73-79. 517

4. Downs, C.C., R.G. White, and B.B. Shepard. 1997. Age at Sexual Maturity, Sex

Ratio, Fecundity, and Longevity of Isolated Headwater Populations of Westslope

Cutthroat Trout. North American Journal of Fisheries Management, 17:1(85-92).

5. Hickman, T., and R.F. Raleigh. 1982. Habitat sustainability index models:

Cutthroat trout. U.S.D.I. Fish and Wildlife Service. FWS/OBS-82/10.5. 38 pp.

6. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

518

Rainbow Trout, Oncorhynchus mykiss (Walbaum, 1792)

Etymology and Synonyms: Oncorhynchus = Greek for “hooked snout”, referring to the elongated hooked jaw, or kype, of a spawning male; mykiss = likely derived from mikizha or mykyz, a native Kamchatkan name for trout.

Description: Body slender, elongated, and slightly laterally compressed. Dorsally dark olive to blue-green; laterally silver to light green with distinct bright pink lateral stripe; ventrally white to silver; heavily spotted with small black spots on head, body, and fins; dorsal fin may have cream colored tip in some populations; pelvic and anal fins generally have a white distal end. Head short. Snout rounded. Eye moderately large. Mouth large, terminal. Frenum absent. Teeth small; present on jaws and tongue. Gill rakers 16-22.

Dorsal fin with 10-12 rays and nearly straight distal end. Adipose fin present. Caudal peduncle stout, thick. Caudal fin moderately forked. Anal fin with 8-12 rays, relatively large. Pelvic fins abdominal with 9-10 rays; insertion slightly posterior to insertion of dorsal fin; small axillary process present. Pectoral fins with 11-17 rays. Lateral line complete with 100-150 small cycloid scales in series. Juveniles with 5-10 dark, vertical bars or “parr marks” present on lateral sides. Spawning males develop a hook, or “kype” on lower jaw, lack breeding tubercles and develop an intensely red lateral stripe.

Similar Species: May be confused with Brown Trout, Brook Trout or Cutthroat Trout.

Rainbow Trout are easily distinguishable from these other trout species by the bright pink lateral stripe. Brown Trout generally lack spots on the caudal fin and display both black and red spots on the body that are outlined in light blue. Brook Trout lack a pink lateral stripe, and instead have well defined red or pink spots outlined in blue. Cutthroat Trout have red to orange slash marks on each underside of the lower jaw. 519

Distribution and Habitat: Native to the Pacific Coast drainages of northeastern Asia,

Canada, and the United States from Alaska to California.2 Introduced widely outside of its native range for sport fishing, including the Dakotas where Rainbow Trout are stocked and maintained in impoundments, tailwaters of the Missouri River reservoirs, and Black

Hills streams. Due to the species being very adaptable to a wide diversity of habitats,

Rainbow Trout have three main ecological forms which are anadromous, (referred to as

“steelhead”), stream resident, and lake and reservoir dwelling. Rainbow Trout strive in clear, well oxygenated waters with optimal temperatures near 12-18 °C (53.6-64.4 °F).1

The species is often found over gravel and rocky substrates, near riffles and areas rich in vegetative cover.1

Reproduction: Self-sustaining populations of Rainbow Trout are known in the Black

Hills of South Dakota, however most other populations throughout the Dakotas require routine stocking, due to conditions which do not allow for natural reproduction. In self- sustaining populations, spawning begins in early spring or summer or when water temperatures reach 6-7 °C (43-45 °F) over gravel substrates.3,5 Unlike steelhead, stream resident and lake and reservoir dwelling Rainbow Trout migrate short distances to spawning grounds that are generally in smaller inlets and tributary streams of lakes and rivers.4 Age at sexual maturity is generally age 2-3, with males often reaching maturity prior to females.1 Female Rainbow Trout prepare a nest, or a “redd”, in gravel substrate often downstream of a pool or near the head of a series of riffles.1,5 Once the eggs are fertilized, the females cover the eggs with gravel. Hatching occurs within 3-4 weeks in water 10-15 °C (50-59 °F).6 520

Age and Growth: Hatchery raised Rainbow Trout are known to exhibit faster growth patterns than wild Rainbow Trout.7,8 Growth varies within populations due to numerous factors such as temperature and diet.7,9,10 Mean lengths-at-age of Rainbow Trout from

Deerfield Reservoir in South Dakota are reported as: age-0, 40 mm (1.57 in); age-1, 60 mm (2.36 in); age-2, 125 mm (4.92 in); age-3, 175 mm (6.88 in); age-4, 210 mm (8.27 in)

TL.11 Capable of reaching 678 mm (26.7 in) TL in the Dakotas. Few Rainbow Trout live past age-6; however, longevity is reported as age-11.12

Food and Feeding: Adult Rainbow Trout feed on a variety of prey items including juvenile and adult forms of aquatic and terrestrial insects, small crustaceans such as zooplankton and scuds, leeches, and fish eggs. In Lake Oahe, Rainbow Trout undergo an ontogenetic diet shift similar to Rainbow Trout in other populations, where they may consume smaller fishes depending on their size and habitat.13,14 Smaller individuals feed predominately on zooplankton and aquatic macroinvertebrates such as mayflies, caddisflies, and stoneflies, especially in the spring and summer months.13,15

Literature Cited:

1. Baxter, G.T., and M.D. Stone. 1995. Fishes of Wyoming. Wyoming Game and

Fish Department. Cheyenne, Wyoming.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Behnke, R.J. 1979. Monograph of the native trouts of the genus Salmo of western

North America. U.S. Fish and Wildlife Service, Region 6, Denver, Colorado.

215pp.

4. Behnke, R.J. 2002. Trout and salmon of North America. The Free Press, Simon

and Schuster, Inc., New York. 521

5. Bjornn, T.C., and D.W. Reiser. 1991. Habitat requirements of salmonids in

streams. Influences of forest and rangeland management on salmonid fishes and

their habitats Pages 83-138 in D. Meehan, editor. American Fisheries Society,

Special Publication 19, Bethesda, Maryland.

6. Davis, J.L. 2012. Contribution of natural recruitment to the rainbow trout

Oncorhynchus mykiss sport fishery in Deerfield Reservoir. Master’s thesis. South

Dakota State University, Brookings, South Dakota.

7. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

8. Kientz, J.L. 2016. Survival, abundance, and relative predation of wild rainbow

trout in the Deerfield Reservoir System, South Dakota. Master’s thesis. South

Dakota State University, Brookings, South Dakota.

9. Koth, R.M. 1980. Food habits and growth of rainbow trout in a prairie pond.

Master’s thesis. South Dakota State University, Brookings, South Dakota.

10. Lynott, S.T., S.D. Bryan, T.D. Hill, and W.G. Duffy. 1995. Monthly and size-

related changes in the diet of rainbow trout in Lake Oahe, South Dakota. Journal

of Freshwater Ecology 10(4): 399-40.

11. Marcogliese, L.A., and J.M. Casselman. 1998. Scale methods for discriminating

between Great Lakes stocks of wild and hatchery rainbow trout, with a measure of

natural recruitment in Lake Ontario. North American Journal of Fisheries

Management 18:253-268. 522

12. Nelitz, M.A., E.A. MacIsaac, and R.M. Peterman. 2007. A science-based

approach for identifying temperature-sensitive streams for rainbow trout. North

American Journal of Fisheries Management 27:405-424.

13. Railsback, S.F., and K.A. Rose. 1999. Bioenergetics modeling of stream trout

growth: temperature and food consumption effects. Transactions of the American

Fisheries Society 128:241-256.

14. Raleigh, R.F., T. Hickman, R.C. Solomon, and P.C. Nelson. 1984. Habitat

suitability information: Rainbow Trout. US Fish and Wildlife Service. FWS/OBS-

82/10.60. 64pp.

15. Yard, M.D., L.G. Coggins Jr., C.V. Baxter, G.E. Bennett, and J. Korman. 2011.

Trout piscivory in the Colorado River, Grand Canyon: effects of turbidity,

temperature, and fish prey availability. Transactions of the American Fisheries

Society 140:471-486.

523

Chinook Salmon, Oncorhynchus tshawytscha (Walbaum, 1792)

Etymology and Synonyms: Oncorhynchus = Greek for “hooked snout”, referring to the elongated hooked jaw, or kype, of a spawning male; tshawytscha = vernacular name of the species in Kamchatka. The common name derives from the Chinook Indians, a northwest Native American tribe who heavily relied on the species.

Description: Body fusiform, laterally compressed. Dorsally dark gray to blue-green; laterally silver to silvery olive-blue; ventrally silver to white; few small dark spots present on dorsal and lateral sides as well as fins; gum of lower jaw black; tongue gray.

Head and snout elongate. Eye moderate, placed laterally on head. Mouth large, terminal, slightly oblique; upper jaw extending past posterior edge of eye. Teeth large, sharp; present on both jaws. Gill rakers 16-26. Dorsal fin with 10-14 rays. Adipose fin present.

Caudal peduncle short and thick. Caudal fin slightly forked. Anal fin with 14-19 rays.

Pelvic fin with 10-11 rays. Pectoral fin with 14-17 rays. Lateral line with 130-165 cycloid scales in series. Spawning males develop a hooked lower jaw, or a “kype”. Both sexes generally darker in color when spawning. Juveniles with 6-12 dark vertical bars or “parr marks” on lateral sides.

Similar Species: Chinook salmon differ from other salmonid species in the Dakotas by having black gums on lower jaw and a gray tongue.

Distribution and Habitat: Native to the Arctic and Pacific drainages in northwestern

North America from Alaska in the north, south to California. Only occur within the

Missouri River and its reservoirs in the Dakotas and Montana. Inhabit cold, clear, well- oxygenated pelagic waters. The closing of the Oahe Dam in 1963 created a coldwater habitat suitable for a salmon fishery in Lake Oahe.8 By 1979, Chinook Salmon 524 successfully migrated their way downstream to Lake Oahe from Lake Sakakawea in

North Dakota where they were initially stocked in 1976.11,13,14 They were first stocked into Lake Oahe in 1982 to diversify the existing fishery.14 As the Missouri River reservoirs warm and begin to thermally stratify, Chinook Salmon migrate deeper into the water column.1,11 Preferred water temperature of adults 11 °C (51.8 °F).9 Preferred water temperature of juveniles 14 °C (57.2 °F).7 Lake Oahe Chinook salmon exhibit poor homing ability.12

Reproduction: Natural reproduction does not take place in the Missouri River due to limited spawning substrate and nonexistent overwinter egg incubation habitat.13

Therefore, the Lake Oahe Chinook Salmon population is maintained by artificial spawning and stocking efforts.12 From October to early November, Chinook salmon are artificially spawned from 2-3 kg (4.41-6.61 lbs.) wild salmon ages 2-4 that ascend fish ladders at Whitlock Bay Spawning Station in Gettysburg, South Dakota.2,12 This size at spawning is considerably less than Chinook salmon in their native range, likely due to differences in their diets.3 Majority of sexually mature individuals artificially spawned in

Lake Oahe are age-3.13 Fecundity in Lake Oahe is generally less than in the species native range.3 Relative fecundity of Lake Oahe females is 3.61 eggs/mm (91.69 eggs/in), and mean total fecundity is 2,708 eggs from a mean female length of 660 mm (25.98 in)

TL.3,6 Eggs large, typically pea-sized. Egg size from Lake Oahe is not related to total length of female.3 Egg survival from Lake Oahe is typically poor during hatchery incubation, and survival varies dramatically depending on the female.3,4 In its native range, Chinook salmon are anadromous, migrating from saltwater (where they spend an average of 3-4 years) to their natal freshwater rivers and streams to spawn over clean 525 gravel substrate. Females deposit eggs within a nest, or redd, which they construct using their body and caudal fin. Shortly after the eggs have been fertilized, females guard and cover them with gravel and then move upstream to continue the spawning act several different times. Semelparous species. Shortly after the spawning season, adults die, and eggs hatch in the spring.

Age and Growth: In Lake Oahe, mean length-at-age of males is slightly larger than females.11 Greatest growth increment occurs from age 1-2.11 Mean lengths-at-age for males of the 1992-1994 year classes from Lake Oahe in 1995 are reported as: age-1, 373 mm (14.69 in) TL; age-2, 572 mm (22.52 in) TL; age-3, 672 mm (26.46 in) TL.11

Chinook salmon age 1-2 growth is reported to be faster than age 1-2 individuals from

Lake Sakakawea.1,11 Few individuals live past age-3 in Lake Oahe, but are capable of reaching age-5 in Lake Sakakawea.1,11,13 Capable of reaching 914.4 mm (36 in) TL and

9.98 kg (22 lbs) in Lake Oahe, and capable of exceeding 45.36 kg (100 lbs.) in its native range. Longevity 9 years.5

Food and Feeding: In Lake Oahe, Chinook salmon are heavily dependent on the abundance of Rainbow Smelt.11 Juveniles consume the greatest amount of diversity in the diets consuming zooplankton, aquatic and terrestrial insects, and both juvenile and adult

Rainbow Smelt.11 Following stocking around May, juveniles immediately feed on invertebrates and within roughly two months switch to preying upon Rainbow Smelt.11

Age-1 individuals rely on age-0 and adult Rainbow Smelt as their primary prey, and age-

3 Chinook Salmon only feed on adult Rainbow Smelt.11 Length of Rainbow Smelt consumed increases with increased Chinook Salmon length.11 In years with relatively low

Rainbow Smelt abundance, diets of adult Chinook salmon include a variety of aquatic 526 invertebrates, zooplankton, and a lesser amount of Rainbow Smelt.3 In their native range,

Chinook salmon consume fish, euphausiids, crab larva, squid and pelagic amphipods.10

Literature Cited:

1. Aadland, L.P.E. 1987. Food habits, distribution, age and growth of chinook

salmon, and predation on newly stocked chinook salmon in Lake Sakakawea,

North Dakota. Ph.D. Dissertation, University of North Dakota, Grand Forks.

2. Barnes, M.E. 2007. Fish hatcheries and stocking practices: past and present. Pages

267-293 in C. Berry, K. Higgins, D. Willis, and S. Chipps, editors. History of

fisheries and fishing in South Dakota. South Dakota Game, Fish and Parks, Pierre.

3. Barnes, M.E., R.P. Hanten, R.J. Cordes, W.A. Sayler, and J. Carreiro. 2000a.

Reproductive performance of inland fall chinook salmon. North American Journal

of Aquaculture 62:203-211.

4. Barnes, M.E., W.A. Sayler, R.J. Cordes, and R.P. Hanten. 2003. Potential

indicators of egg viability in landlocked fall chinook salmon spawn with or

without the presence of overripe eggs. North American Journal of Aquaculture

65:49-55.

5. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

6. Becket, K.H., M.E. Barnes, D.J. Durben, and T.M. Parker. 2015. Landlocked fall

chinook salmon ovarian fluid turbidity and egg survival. North American Journal

of Aquaculture 77:18-21.

7. Brett, J.R. 1952. Temperature tolerance in young Pacific salmon, genus

Oncorhynchus. Journal of the Fisheries Research Board of Canada 9:265-323. 527

8. Cordes, R. 2007. Cold-water fish species. Pages 201-211 in C. Berry, K. Higgins,

D. Willis, and S. Chipps, editors. History of fisheries and fishing in South Dakota.

South Dakota Game, Fish and Parks, Pierre.

9. Haynes, J.M., and C.J. Keleher. 1986. Movements of Pacific salmon in Lake

Ontario in spring and summer: evidence of wide spatial dispersal. Journal of

Freshwater Ecology 3:289-297.

10. Higgs, D.A., J.S. Macdonald, C.D. Levings, and B.S. Dosanjh. 1995. Nutrition

and feeding habits in relation to life history stage. Pages 159-316 in C. Grott, L.

Margolis, and W.C. Clarke, editors. Physiological ecology of Pacific salmon.

University of British Columbia Press, Vancouver.

11. Hill, T.D. 1997. Life history and bioenergetics of chinook salmon in Lake Oahe,

South Dakota. Ph.D. Dissertation, South Dakota State University, Brookings.

12. Lott, J., G. Marrone, and D. Stout. 1997. Influence of size-and-date at stocking,

imprinting attempts and growth on initial survival, homing ability, maturation

patterns and angler harvest of chinook salmon in Lake Oahe, South Dakota. South

Dakota Department of Game, Fish and Parks, Progress Report 97-20, Pierre.

13. Marrone, G.M., and D.A. Stout. 1996. Whitlocks Bay spawning and imprinting

station annual report 1995. Wildlife Division, South Dakota Department of Game,

Fish and Parks, Pierre.

14. Warnick, D.C. 1987. The introduction of selected fish species into the Missouri

River system, 1987-1985. Wildlife Division, South Dakota Department of Game,

Fish and Parks, Progress Report 87-10, Pierre.

528

Brown Trout, Salmo trutta (Linnaeus, 1758)

Etymology and Synonyms: Salmo = Latin for “Atlantic salmon”; trutta = Latin name for “trout”.

Description: Body slender, elongate, slightly laterally compressed. Dorsally brown to olive with large dark brown to black spots which may be as large as pupil, outlined in tan to light blue, leading onto lateral sides; laterally light olive to tan with continued large, round to irregularly shaped dark spots outlined in light tan or blue, as well as less numerous small, prominent red spots also outlined in light tan or blue; ventrally cream to white; dorsal fin with numerous small dark brown spots; adipose fin with orange to red coloration on distal end; caudal fin with little to no spots; anal fin with white edge on distal end. Head small. Snout rounded. Eye moderately large, placed laterally on head.

Mouth large, terminal, slightly oblique; upper jaw extending to or slightly beyond posterior edge of eye. Teeth present on both jaws, tongue, vomer and palatines. Gill rakers 14-20. Dorsal fin with 12-14 rays. Adipose fin present. Caudal peduncle stout.

Caudal fin square to slightly forked. Anal fin with 10-12 rays. Pelvic fins abdominal with

9-10 rays; small axillary process present. Pectoral fins with 13-14 rays. Lateral line complete with 120-130 cycloid scales in series. Spawning larger males develop a hooked lower jaw, or “kype”. Juveniles with less numerous spots on lateral sides and 7-11 dark, oval shaped blotches, or “parr marks” on lateral sides.

Similar Species: Brook Trout with well-defined small pink or red spots outlined in blue against a dark olive background on lateral sides, and with well-defined black stripes and white edging on distal ends on the lower lobe of the caudal, anal, pelvic, and pectoral fins. Cutthroat Trout display numerous small black spots on body extending onto the 529 caudal fin, and have reddish-orange slash marks on each underside of the lower jaw.

Rainbow Trout with a bright pink to reddish lateral stripe present and numerous small black spots on body.

Distribution and Habitat: Native to Europe, northern Africa and western Asia.21 Widely introduced outside its native range due to sportfishing, and now have a world-wide distribution.10 Introduced into the United States in 1883.4 Mainly occur west of the

Missouri River in the Dakotas. Adults inhabit deeper areas with little flow in streams or rivers with vegetative cover and gravel or stony substrate.19 In the Black Hills of South

Dakota, runs and pools are used more often than riffles in fall and winter, and runs are used more often than pools or riffles in the spring.12 A rather sedentary species with small home ranges and migration movements.2,12 Females typically more migratory than males.7,10,19 Changes in water temperature and flow induce up- and downstream migration movements.17,19 Individuals from the Black Hills migrated an average of 23 m

(75.46 ft.) annually and ranged 2-150 m (6.56-492.13 ft.) annually with the greatest amount of movement during fall and spring.12 Activity levels decrease and migration to deeper overwintering habitat occurs when water temperatures drops below 8°C

(46.4°F).11,12,19 Fry often move downstream post hatching and are associated with shallow, slow-flowing areas <20-30 cm (7.87-11.81 in) in depth near shorelines or banks of their natal streams with water velocity 0-20 cm/s and sand or gravel substrate.1,9,19,22

Reproduction: Spawning migrations begin in the fall. Individuals from the Black Hills of South Dakota migrate up to 1,091 m (0.68 mi) upstream to spawn, and often return within a few meters of their prespawning localities.12 Spawning takes place in fall or early winter when water temperatures reach 3-13°C (37.4-55.4°F) in areas with water 530 velocity 10.8-80.2 cm/s at depths roughly 25.5 cm (10.04 in) in gravel substrate near instream cover.23 Sexual maturity variable and can be reached at age 1-10, with males maturing earlier than females.19 Maturation occurs later in cold localities such as northern rivers and mountain lakes.16,19 Several males court a single female over the nest, but only one large male may fertilize the majority of the eggs.15,19,20 Females construct nests, or redds, within gravel or sand substrate that are a minimum of 14 cm (5.51 in) deep.23

Fecundity dependent on size and age of female. Fractional spawners, with females depositing only a portion of their eggs each time.19 After eggs are deposited, females cover the eggs with stones and gravel where they will remain for several months.

Females then abandon the nest and leave the male to continue spawning with other females.19 Egg size increases with the size of the female.19 Hatching takes place in spring.

Egg mortality is low (<5%) at 9-10°C (48.2-50°F), but increases with rising temperatures

(50% at 12°C (53.6°F) and 100% at temperatures >15.5°C (59.9°F)).1,8,18

Age and Growth: Growth highly variable among populations due to variables such as temperature and food availability.19 Growth known to be density dependent in both streams and lakes.13,14,19 Larvae roughly 20 mm (0.79 in) TL at hatching, and size may decrease with an increased incubation temperature.19 Optimal growth temperature reported as 13-18°C (55.4-64.4°F).19 Upper critical temperature for growth is 25-26°C

(77-78.8°F), and the lower critical temperature for growth is 3-6°C (37.4-42.8°F).19

Medium- to large-sized fish often reaching 254-381 mm (10-15 in) TL. Capable of reaching 762 mm (30 in) TL in the Dakotas. Longevity 20 years.6,19

Food and Feeding: Opportunistic, voracious, visual predator. Larvae and juveniles consume aquatic insect larvae such as chironomids near the shorelines and surface.19 Diet 531 varies with season, habitat, age, and size of fish.5,19 Size of prey increases with size of fish. Adults primarily feed during the day on aquatic invertebrates and larvae such as

Ephemeroptera, Plecoptera, and Trichoptera, near the surface and shorelines. Adults also consume terrestrial insects, worms, crayfish, and small fish. In lakes, zoobenthos and zooplankton are an important food source.19 Optimum feeding temperature 15.6°C

(60.08°F).3

Literature Cited:

1. Armstrong, J.D., P.S. Kemp, G.J.A. Kennedy, M. Ladle, and N.J. Milner. 2003.

Habitat requirements of Atlantic salmon and brown trout in rivers and streams.

Fisheries Research 62:143-170.

2. Bachman, R.A. 1984. Foraging behavior of free-ranging wild and hatchery brown

trout in a stream. Transactions of the American Fisheries Society 113:1-32.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Behnke, R.J. 2002. Trout and salmon of North America. The Free Press, Simon

and Schuster, Inc., New York.

5. Bridcut, E.E., and P.S. Giller. 1995. Diet variability and foraging strategies on

brown trout (Salmo trutta): an analysis from subpopulations to individuals.

Canadian Journal of Fisheries and Aquatic Sciences 52:2543-2552.

6. Brown, M.E. 1957. Experimental studies on growth. Pages 361-400 in The

Physiology of Fishes. Vol. 1, M.E. Brown (editor). Academic Press, New York.

7. Campbell, J.S. 1977. Spawning characteristics of brown trout and sea trout Salmo

trutta L. in Kirk Burn, River Tweed, Scotland. Journal of Fish Biology 11:217-

229. 532

8. Crisp, D.T. 1993. The environmental requirements of salmon and trout in fresh

water. Freshwater Forum 3:176-202.

9. Elliott, J.M. 1986. Spatial distribution and behavioural movements of migratory

trout Salmo trutta in a Lake District stream. Journal of Animal Ecology 55:907-

922.

10. Elliott, J.M. 1994. Quantitative Ecology and the Brown Trout. Oxford: Oxford

University Press.

11. Heggenes, J. and J.G. Dokk. 2001. Contrasting temperatures, waterflows, and

light: seasonal habitat selection by young Atlantic salmon and brown trout in a

boreonemoral river. Regulated Rivers: Research and Management 17:623-635.

12. James, D.A., J.W. Erickson, and B.A. Barton. 2007. Brown trout seasonal

movement patterns and habitat use in an urbanized South Dakota stream. North

American Journal of Fisheries Management 27:978-985.

13. Jenkins, T.M., S. Diehl, K.W. Kratz, and S.D. Cooper. 1999. Effects of

population density on individual growth of brown trout in streams. Ecology

80:941-956.

14. Jensen, K.W. 1977. On the dynamics and exploitation of the population of brown

trout, Salmo trutta L., in Lake Øvre Heimdalsvatn, southern Norway. Report of

Institute of Freshwater Research, Drottningholm 56:18-69.

15. Jones, J.W. and J.N. Ball. 1954. The spawning behavior of brown trout and

salmon. British Journal of Animal Behaviour 2:103-114. 533

16. Jonsson, B., J.H. L’Abèe-Lund, T.G. Heggberget, A.J. Jensen, B.O. Johnsen, T.F.

Nӕsje, and L.M. Sӕttem. 1991c. Longevity, body size and growth in anadromous

brown trout. Canadian Journal of Fisheries and Aquatic Sciences 48:1838-1845.

17. Jonsson, N. 1991. Influence of water flow, water temperature and light on fish

migration in rivers. Nordic Journal of Freshwater Research 66:20-35.

18. Jowett, I.G. 1992. Models of abundance of large brown trout in New Zealand

rivers. North American Journal of Fish Management 12:417-432.

19. Klemetsen, A., P.-A., Amundsen, J.B. Dempson, B. Jonsson, N. Jonsson, M.F.

O’connell, and E. Mortensen. 2003. Atlantic salmon Salmo salar L., brown trout

Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of

their life histories. Ecology of Freshwater Fish 12:1-59.

20. Largiander, C.R., A. Estoup, F. Lecerf, A. Champigneulle, and R. Guyomard.

2001. Microsatellite analysis of polyandry and spawning site competition in

brown trout (Salmo trutta L.). Genetics Selection Evolution 33:205-222.

21. MacCrimmon, H.R., T.L. Marshall, and B.L. Gotos. 1970. World distribution of

brown trout, Salmo trutta: further observations. Journal of the Fisheries Research

Board of Canada 27:811-818.

22. Roussel, J.M., and A. Bardonnet. 1999. Ontogeny of diel pattern of stream-margin

habitat use by emerging brown trout, Salmo trutta, in experimental channels:

influence of food and predator presence. Environmental Biology of Fishes

56:253-262. 534

23. Witzel, L. and H. MacCrimmon. 1983. Redd-site selection by brook trout and

brown trout in southwestern Ontario streams. Transactions of the American

Fisheries Society 112:760-771.

535

Brook Trout, Salvelinus fontinalis (Mitchill, 1814)

Etymology and Synonyms: Salvelinus = a deviation of the old vernacular name for the

“char”; fontinalis = “of springs”, likely in reference to the species habitat.

Description: Body slender, elongate, slightly laterally compressed. Dorsally olive to dark bluish-gray with irregular light gray blotches, or “vermiculations”; lateral sides with light, well-defined red or pink spots outlined in light blue against a dark olive background with light cream to gray irregular blotches; ventrally light yellow, cream to white; dorsal and caudal fin with bark brown to black spots; lower lobe of caudal, anal, pelvic, and pectoral fins with light red to orange coloring leading into a distinct black stripe followed by a white edge on distal end. Head small. Snout rounded. Eye moderately large, placed laterally on head. Mouth large, terminal, slightly oblique; upper jaw extends beyond posterior edge of eye. Teeth present on both jaws and tongue. Gill rakers 9-12. Dorsal fin with 10-14 rays. Adipose fin present. Caudal peduncle stout. Caudal fin square shaped to slightly forked. Anal fin with 9-14 rays. Pelvic fins abdominal with 8-10 rays; small axillary process present. Pectoral fins with 11-14 rays. Lateral line complete with 210-

240 cycloid scales in series. Spawning adults develop black gum lines on the mouth and bright coloration, especially males. Lateral spots become more intense, and the lateral- ventral sides develop a bright gold to red color with black pigmented striped on each side of the belly. Pectoral, pelvic, and anal fins also develop bright red coloration. Large males may develop a hooked lower jaw, or “kype”. Juveniles darker on dorsal sides with

7-9 dark, oval shaped blotches, or “parr marks” on lateral sides.

Similar Species: Distinguishable from other species in the Dakotas by the well-defined pink or red spots outlined in blue against a dark olive background on lateral sides, as well 536 as the black stripes present on the lower lobe of the caudal, anal, pelvic, and pectoral fins.

Brown Trout with black to brown spots outlined in blue on light olive to tan lateral sides, and generally lack numerous dark spots on caudal fin. Rainbow Trout with a pinkish red lateral stripe present and numerous small black spots. Lake Trout have a slate gray to emerald green head and body that is heavily marked with creamy white spots, and also have a forked caudal fin.

Distribution and Habitat: Native to most of eastern Canada and the northeastern United

States from eastern Minnesota and the upper Mississippi River basin in the west, east throughout the Great Lakes, Hudson Bay and Atlantic drainages from New York, south throughout the Appalachian Mountains to northern Georgia. Introduced outside its native range to much of the western United States due to sport fishing. Majority of Brook Trout in the Dakotas occur in the Black Hills of South Dakota, but have also reported in the

Missouri River basin, the Red River basin in eastern North Dakota, and the Minnesota

River drainage of eastern South Dakota. Inhabit cool, clear, spring-fed ground waters of headwater streams, rivers, and oligotrophic lakes with gravel substrate and aquatic vegetation or submerged structure.12 Migrate downstream post spawning to deeper, quiet waters, sometimes within eddies and under banks in winter.4 A rather sedentary species, with little movement annually besides pre- and post-spawning. Preferred water temperature 11-16°C (51.8-60.8°F).9 Fry have an optimum water temperature of 8-12°C

(46.4-53.6°F), and inhabit shallow areas near shorelines with aquatic vegetation used as cover.15,16 Sensitive to moderate turbidity and warm water temperatures.

Reproduction: Migrate upstream into headwater streams or tributaries during the late summer and fall. Males reach sexual maturity earlier than females, and age at maturation 537 generally early in life but variable with populations.16 Spawning takes place late fall when water temperatures reach roughly 4-10°C (39.2-50.0°F).16 Females spend up to two days constructing nests using their body and caudal fin to sweep depressions into coarse or gravel substrate in shallow areas roughly 36.-61 cm (1.18-2.00 ft.) in depth and surface water velocities of roughly 13-24 cm/s.5,10,11 Females prefer to spawn with larger males, and may delay spawning when paired with smaller males.6 Fecundity increases with size and age of female. In Wisconsin, a 127 mm (5.0 in) TL female produced <100 eggs, and a 356 mm (14.02 in) TL female produced roughly 1,200 eggs.3 After eggs are deposited, females stir up gravel to cover the eggs before they desert the nest and leave the male to provide parental care for up to two weeks.11 Hatching takes place late winter to early spring depending on water temperature. Hatching occurs within 45 days at 10°C (50.0°F) and 165 days at 2.8°C (37.04°F).7,16

Age and Growth: Optimal water temperature for growth and survival of juveniles is 12-

15°C (53.6-59°F).14 Growth rates fastest in spring and summer.12 Optimal growth conditions for adults occur in water 9-17°C (48.2-62.6°F).2,16 Growth rates known to be higher and greater lengths are achieved in areas with low Brown Trout abundance.8,12

Annual growth rates may differ from variability in water temperature, species density, prey availability.12 Density-dependent growth often occurs with high water temperatures and limited prey resources.17,18 Male growth rates faster than females. Capable of exceeding 508 mm (20 in) TL. Average longevity in streams 3-4 years, but capable of reaching 14 years in high elevation streams and rivers.13

Food and Feeding: Opportunistic visual feeder. Proportion of diet items depends upon availability. Feeding takes place during the day on a wide variety of aquatic 538 macroinvertebrates such as Diptera, Trichoptera, and Ephemeroptera, which make up an important part of the diet.1 Terrestrial insects, crustaceans, and annelids also generally consumed. Larger adults may occasionally feed on smaller fish or eggs, sometimes of its own kind.

Literature Cited:

1. Allan, J.D. 1981. Determinants of diet of brook trout (Salvelinus fontinalis) in a

mountain stream. Canadian Journal of Fisheries and Aquatic Sciences 38:184-

192.

2. Baldwin, N.S. 1956. Food consumption and growth of brook trout at different

temperatures. Transactions of the American Fisheries Society 86:323-328.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Benson, N.G. 1955. Observation on anchor ice in a Michigan trout stream.

Ecology 36:529-530.

5. Bernier-Borugault, I., and P. Magnan. 2002. Factors affecting red site selection,

hatching, and emergence of brook charr, Salvelinus fontinalis, in an artificially

enhanced site. Environmental Biology of Fishes 64:333-341.

6. Blanchfield, P.J., and M.S. Ridgway. 1999. The cost of peripheral males in a

brook trout mating system. Animal Behavior 57:537-544.

7. Brasch, J., J. McFadden, and S. Kmiotek. 1958. Brook trout. Life history,

ecology, and management. Wisconsin Department of Natural Resources

Publication 226. 539

8. Carlson, S.M., A.P. Hendry, and B.H. Letcher. 2007. Growth rate differences

between resident native brook trout and non-native brown trout. Journal of Fish

Biology 71:1430-1447.

9. Coutant, C.C. 1977. Compilation of temperature preference data. Journal of the

Fisheries Research Board of Canada 34:739-745.

10. Essington, T.E., P.W. Sorenson, and D.G. Paron. 1998. High rates of red

superimposition by brook trout (Salvelinus fontinalis) and brown trout (Salmo

trutta) in a Minnesota stream cannot be explained by habitat availability alone.

Canadian Journal of Fisheries and Aquatic Sciences 55:2310-2316.

11. Hazzard, A.S. 1932. Some phases of the life history of the eastern brook trout

Salvelinus fontinalus Mitchill. Transactions of the American Fisheries Society

62:344-350.

12. Hoxmeier, R.J.H., and D.J. Dieterman. 2013. Seasonal movement, growth and

survival of brook trout in sympatry with brown trout in Midwestern US streams.

Ecology of Freshwater Fish 22:530-542.

13. Kennedy, B.M., D.P. Peterson, and K.D. Fausch. 2003. Different life histories of

brook trout populations invading mid-elevation and high-elevation cutthroat trout

streams in Colorado. Western North American Naturalist 63:215-223.

14. McCormick, J.H., K.E.F. Hokanson, and B.R. Jones. 1972. Effects of temperature

on growth and survival of young brook trout, Salvelinus fontinalis. Journal of the

Fisheries Research Board of Canada 29:1107-1112. 540

15. Peterson, R.H., A.M. Sutterlin, and J.L. Metcalfe. 1979. Temperature preference

of several species of Salmo and Salvelinus and some of their hybrids. Journal of

the Fisheries Research Board of Canada 36:1137-1140.

16. Raleigh, R.F. 1982. Habitat suitability index models: brook trout. United States

Fish and Wildlife Service FWS/OBS-82/10.24.

17. Utz, R.M., and K.J. Hartman. 2009. Density-dependent individual growth and size

dynamics of central Appalachian brook trout (Salvelinus fontinalis). Canadian

Journal of Fisheries and Aquatic Sciences 66:1072-1080.

18. Xu, C.L., B.H. Letcher, and K.H. Nislow. 2010. Size-dependent survival of brook

trout Salvelinus fontinalis in summer: effects of water temperature and stream

flow. Journal of Fish Biology 76:2342-2369.

541

Lake Trout, Salvelinus namaycush (Walbaum, 1792)

Etymology and Synonyms: Salvelinus = old name for char; namaycush = old American

Indian name.

Description: Body elongate, moderately slender. Body slate gray to dark emerald green in color with creamy white spots present on head, body, and fins, fading to a faint white ventral side; pectoral, pelvic, anal, and caudal fins can display slight orange tint; pectoral, pelvic and anal fins with white bordering on distal ends. Juveniles with thick, dark, vertical parr marks on lateral sides. Head short. Snout moderately conical, slightly protrudes beyond upper jaw. Eye small. Mouth large, terminal, maxillary extends well beyond eye. Teeth present on upper and lower jaws; point posterior on prevomer.

Mandibular pores 9-10. Gill rakers 14-25. Branchiostegal rays 10-13. Dorsal fin rays 9-

14. Adipose fin present. Caudal peduncle slender. Caudal fin deeply forked. Anal fin rays

8-11. Pelvic fin rays 8-11. Pectoral fin rays 13-17. Lateral line complete with 170-220.

Scales small, cycloid, deeply embedded. Spawning males with dark lateral stripe.

Similar Species: Brook Trout have a much more weakly forked caudal fin, and display red spots outlined in blue on their lateral sides, whereas Lake Trout lack red spots, and instead have numerous creamy white spots present on their head, body, and fins. Splake

Trout (a hybrid resulting from the crossing of two fish species, a female Lake Trout and a male Brook Trout) occur sympatrically with Lake Trout populations in the Dakotas and may also be misidentified. Splake Trout appearance is astonishingly similar to Lake

Trout, however the caudal fin is not as deeply forked as in Lake Trout.

Distribution and Habitat: Native range includes most of Canada, and the northern part of the United States from Montana to the Great Lakes, and east to Maine. Occurs in Lake 542

Sakakawea in North Dakota from stockings in 1973 and 1974.8 Present in Lake Oahe,

Lake Sharpe, and Lake Pactola, Lake Deerfield, and Pactola Stilling Pond in South

Dakota. Frequently found in well-oxygenated, cold, deeper waters of lakes with rocky substrate. May move throughout the water column during seasons. In the Great Lakes, they are most abundant at depths between 30-90 m (110-300 ft.).1 Preferred water temperature is roughly 10°C (50°F), but can tolerate temperatures up to 25°C (77°F).1,5

Light intensity is known to strongly affect winter habitat selection of Lake Trout, due to foraging efficiency.4

Reproduction: Spawning behavior of Lake Trout is poorly understood compared to other salmonid species. Lake Trout are unique among other salmonids in that spawning primarily takes place in lakes rather than rivers and streams. Spawning begins in the fall, usually from October to late November, and occurs in the late evenings. Lake Trout migrate to shallow waters to spawn over gravel or rock substrate. Most return to the same spawning grounds each year.3 Sexual maturity of males, typically age-4; females age-5.

Males enter the spawning grounds earlier than females.7 As a female enters, one or more males will hover and remain in close contact while courting the female to her desired spawning area.2 The group will then settle on the substrate and quiver their bodies together to fertilize the eggs.2 No nest is prepared and no parental care is given. Eggs sink to the bottom after dispersed, and are 5-6 mm (0.20-0.24 in) in diameter.1 Eggs incubate for 15-21 weeks before hatching in early spring. Fry leave spawning beds and move to deeper water after yolk sac is absorbed.

Age and Growth: Average lifespan 10-20 years. Demonstrate relatively rapid growth within the first five years. Average 381-762 mm (15-30 in) TL in larger, deeper lakes, 543 and 304-406 mm (12-16 in) TL in smaller bodies of water. Capable of reaching lengths of

1 m (3.2 ft.) and exceeding 50 lbs.9 Length-at-age for Lake Trout in Lake Superior were documented as: age-1, 102 mm (4.02 in) TL; age-2, 160 mm (6.30 in) TL; age-3, 216 mm

(8.50 in) TL; age-4, 279 mm (10.98 in) TL; age-5, 351 mm (13.82 in) TL; age-6, 427 mm

(16.81 in) TL; age-7, 500 mm (19.69 in) TL; age-8, 579 mm (22.80 in) TL.1

Food and Feeding: Visual feeders. Lake Trout residing in larger lakes are primarily piscivorous, and feed heavily on pelagic fish, littoral minnows and shiners in winters, and zooplankton in the summers.4 Lake Trout in smaller lakes with less prey fish abundance mainly feed on zooplankton, crustaceans, and aquatic invertebrates. It is known that Lake

Trout who feed primarily on zooplankton and invertebrates mature earlier, grow slower, attain a smaller maximum size, and die more quickly than Lake Trout who primarily consume fish.6 Juveniles feed on crustaceans and aquatic invertebrates.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Binder, T.R., et al. 2014. New insight into the spawning behavior of lake trout,

Salvelinus namaycush, from a recovering population in the Laurentian Great

Lakes.

3. Binder, T.R., et al. 2015. Spawning site fidelity of wild and hatchery lake trout

(Salvelinus namaycush) in northern Lake Huron. Canadian Journal of Fisheries

and Aquatic Sciences, 73(1),18-34.

4. Blanchfield, P.J., Tate, L.S., Plumb, J.M. et al. 2009. Seasonal habitat selection by

lake trout (Salvelinus namaycush) in a small Canadian shield lake: constraints

imposed by winter conditions. Aquatic Ecology, 43:777-787. 544

5. Eschmeyer, P.H. 1957. The lake trout (Salvelinus namaycush). U.S. Fish Wildl.

Serv. Fish. Leaflet No. 441. 11pp.

6. Martin, N.V. 1966. The significance of food habits in the biology, exploitation,

and management of Algonquin park, Ontario, lake trout. Transactions of the

American Fisheries Society 95:415-422.

7. Muir, A.M., et al. 2012b. Lake charr Salvelinus namaycush spawning behavior:

new field observations and a review of current knowledge. Rev Fish Biol Fisher

22:575-593.

8. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

9. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

545

CHAPTER 15

FAMILY OSMERIDAE

Introduction

The Smelt family, Osmeridae, consists of fifteen extant species within seven genera worldwide. Ten species within six genera occur within North American waters.

Most of the species in the family are entirely marine and originate from the North Pacific and Northern Atlantic oceans near North America and Northern Asia where they are a popular commercial fish. However, some species are anadromous, and other species have been introduced to persist in landlocked, freshwater populations, such as the Rainbow

Smelt, Osmerus mordax, which occurs in the Dakotas within the Missouri River reservoirs.

Smelts are characterized by having a slender and streamlined body with pointed snouts, the presence of a small adipose fin positioned anterior to the single dorsal fin, and a deeply forked caudal fin. Most species of smelt are smaller, and only seldom reach more than 203-254 mm (8-10 in) TL. Smelt are often confused with members of the salmon family, Salmonidae, which they are distantly related to. Members of Osmeridae can be distinguished from members of Salmonidae by the lack of an axillary process, which is present at the anterior base of the pelvic fins on salmonids. In our region, the

Rainbow Smelt may also be initially misidentified with species other than salmonids such as the Trout Perch, Percopsis omiscomaysus, due to the small adipose fin. However,

Trout Perch display a complete lateral line, and Rainbow Smelt have an incomplete lateral line. 546

Smelt are pelagic fish and prefer the open and cool waters of the ocean, rivers, lakes, and reservoirs. In the Dakotas, Rainbow Smelt inhabit the deep, dark, cold, stratified waters of the Missouri River reservoirs where they form large schools, often so near dams and intake structures. Only during the spawning season, which occurs shortly after ice-out, will Rainbow Smelt migrate upstream into smaller tributaries to spawn during the night in shallow water with gravel substrate. Members of the Smelt family are carnivorous, and primarily feed on large zooplankton which they devour using their small teeth located on both jaws and the tongue. Adult smelt also consume smaller fishes, larvae, and fry of other fish species, but are known to be cannibalistic, consuming other smaller Rainbow Smelt individuals, larvae and fry as well.

In the Dakotas, Rainbow Smelt are known to be a dominant pelagic prey item for adult Walleye, Sander vitreus, when abundant in the Missouri River reservoirs, and are thought to be a main factor contributing to the fluctuating Walleye population dynamics.

Studies in Lake Oahe, South Dakota, have recently shown that the Rainbow Smelt population significantly decreases during periods of high entrainment caused by increased discharge, which leads to periods of wide fluctuations in Rainbow Smelt abundance. Within this system, Walleye growth rates have been shown to increase exceptionally with a higher abundance of Rainbow Smelt present. In contrary, when abundance of Rainbow Smelt decreases, they are still consumed by adult Walleye, however the condition and growth of the Walleye decreases.

547

Rainbow Smelt, Osmerus mordax (Mitchill, 1814)

Etymology and Synonyms: Osmerus = Greek for osme, meaning “odorous”, referring to the fish’s scent that often reminds one of cucumbers; mordax = biting.

Description: Body fusiform, slender and laterally compressed. Dorsally olive green to dark gray; laterally silver with hues of purple, blue, and/ or light pink iridescence; color tends to fade rather quickly when removed from the water; ventrally silver to white; fins generally clear. Head and snout moderate, tapering to a point. Eyes large, placed laterally on head. Mouth terminal, slightly oblique; relatively large with the upper jaw extending to center of eye or just beyond; lower jaw slightly protrudes past upper jaw. Teeth present on both jaws and tongue. Gill rakers 26-34, long and slender. Dorsal fin positioned approximately equidistant between snout and caudal fin base with 8-11 rays. Adipose fin present; small with nearly rounded distal end. Caudal peduncle relatively short and thick.

Caudal fin forked. Anal fin slightly concave with 11-16 rays. Pelvic fins with 8 rays; insertion distinctly inferior to dorsal fin insertion. Pectoral fins with 11-14 rays. Lateral line absent with 60-72 cycloid scales in series. Breeding males develop small tubercles on head, fins, and body.

Similar Species: May be mistaken for Emerald Shiner, Trout-perch, or other trout species due to the presence of an adipose fin. Although Emerald Shiner may have similar coloration, they lack an adipose fin, have no teeth, and have a complete lateral line.

Trout-perch and other trout species in the Dakotas can be distinguished by their complete lateral lines.

Distribution and Habitat: Native to the northeastern Atlantic and Pacific-Arctic drainages.16 While Rainbow Smelt are generally an anadromous species, they are widely 548 introduced throughout North America in several landlocked habitats such as lakes, reservoirs, and streams as a pelagic forage fish for sport fish predators such as

Walleye.4,7,9 Rainbow Smelt are primarily found throughout the Missouri River and its reservoirs in the Dakotas. The species prefers cooler waters, and in Lake Oahe, Rainbow

Smelt occupy waters 5-14 °C (41-57 °F).4 Hydroacoustic surveys have shown that

Rainbow Smelt avoid light and are found aggregating near the bottom during the day.4,10,15 At night, most Rainbow Smelt are associated with the thermocline, indicating that water temperature acts as a barrier for the species when they migrate up in the water column at night.4,10,15 The species is also known to congregate near dams and intake structures.10,8

Reproduction: Spawning is initiated shortly after ice-out, or when water temperatures hover around 4.4 °C (40 °F).2 Spawning migrations consist of movements made up clear, swift tributary streams, shorelines or shallow areas <0.6 m (2 ft.) with rock or gravel substrate.18 Age at sexual maturity varies depending on the population with the majority spawning at age-1, however other populations have been known to spawn at age 2-4.5,13,14

In some populations, males have been observed maturing one year earlier than females.13

No nest is constructed, and no parental care is given. Fecundity from females 185-224 mm (7.3-8.8 in) TL ranged 21,534-40,894 eggs.2 Eggs 0.9-1.3 mm (0.04-0.05 in) in diameter, demersal and adhesive to submerged vegetation or substrate.2,6 Hatching takes place in 2-3 weeks.19

Age and Growth: Larvae approximately 5.5-6.0 mm (0.22-0.23 in) TL.2 Females generally reach larger sizes and live longer than males.1 Mean female lengths-at-age of

Rainbow Smelt from Wisconsin are reported as: age-1, 66 mm (2.60 in) TL; age-2, 150 549 mm (5.91 in) TL; age-3, 193 mm (7.60 in) TL; age-4, 221 mm (8.70 in) TL; age-5, 239 mm (9.41 in) TL; age-6, 257 mm (10.12 in) TL; age-7, 310 mm (12.20 in) TL.2 Rainbow

Smelt are a rather short-lived species with many rarely surviving past age-3.1 Longevity 8 years.1,11

Food and Feeding: In general, juvenile Rainbow Smelt are planktivorous, and consume large amounts of zooplankton such as Copepods and Cladocerans.6 Adults tend to be cannibalistic and consume the highest number of their own species compared to any other species.3 Adults also consume fry and larvae of other pelagic forage species, as well as larger crustaceans and immature insects.12,17,20

Literature Cited:

1. Bailey, M.M. 1964. Age, growth maturity and sex composition of the American

Smelt, Osmerus mordax (Mitchill), of western Lake Superior. Transactions of the

American Fisheries Society 93:382-395.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Burbidge, R.G. 1969. Age, growth, length-weight relationship, sex ratio, and food

habits of American smelt, Osmerus mordax (Mitchill), from Gull Lake, Michigan.

Transactions of the American Fisheries Society, 98:631-640.

4. Burczynski, J.J., P.H. Michalets, and G.M. Marrone. 1987. Hydroacoustic

assessment of the abundance and distribution of rainbow smelt in Lake Oahe.

North American Journal of Fisheries Management 7:106-116.

5. Enterline, C.L. 2013. Understanding spawning behavior and habitat use by

anadromous rainbow smelt (Osmerus mordax) using passive integrated 550

transponder systems and telemetry. Master’s Theses and Capstones, University of

New Hampshire. 1264.

6. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

7. Fincel, M. J., D. J. Dembkowski, and S. R. Chipps. 2014. Influence of variable

rainbow smelt and gizzard shad abundance on walleye diets and growth. Lake and

Reservoir Management 30:258-267.

8. Fincel, M.J., W.J. Radigan, and C.M. Longhenry. 2016. Entrainment of Rainbow

Smelt through Oahe dam during the 2011 Missouri River flood. North American

Journal of Fisheries Management, 36: 844-851.

9. Graeb, B.D., S.R. Chipps, D.W. Willis, J.P. Lott, R.P. Hanten, W. Nelson-

Stastny, and J.W. Erickson. 2008. Walleye response to Rainbow Smelt population

decline and liberalized angling regulations in a Missouri River reservoir. Pages

275–291 in M. S. Allen, S. Sammons, and M. J. Maceina, editors. Balancing

fisheries management and water uses for impounded river systems, American

Fisheries Society, Symposium 62, Bethesda, Maryland.

10. Hamel, M.J., M.L. Brown, and S.R. Chipps. 2008. Behavioral responses of

Rainbow Smelt to in situ strobe lights. North American Journal of Fisheries

Management, 28:2, 394-401.

11. Kirn, R.A., and G.W. Laber. 1996. Growth and survival of Rainbow Smelt, and

their role as prey for stocked salmonids in Lake Champlain. Transactions of the

American Fisheries Society 125:87096. 551

12. Lantry, B.F., Stewart, D.J., 2000. Population dynamics of rainbow smelt

(Osmerus mordax) in Lakes Ontario and Erie: a modeling analysis of cannibalism

effects. Canadian Journal of Fisheries and Aquatic Sciences 57:1594-1606.

13. McKenzie, R. A. 1958. Age and growth of smelt, Osmerus mordax (Mitchill), of

the Miramichi River, New Brunswick. Journal of the Fisheries Research Board of

Canada 15:1313–1327.

14. Murawski, S. A., and C. F. Cole. 1978. Population dynamics of anadromous

Rainbow Smelt Osmerus mordax, in a Massachusetts river system. Transactions

of the American Fisheries Society 107:535–542.

15. Nelson-Stastny, W. 2001. Estimates of abundance, biomass, and distribution of

rainbow smelt and other pelagic fish in Lake Oahe using hydroacoustic

techniques, 1996-1999. South Dakota Game, Fish and Parks, Annual Report 01-

06, Pierre.

16. O’Brien, T.P., W.W. Taylor, E.F. Roseman, C.P. Madenjian, and S.C. Riley.

2014. Ecological factors affecting rainbow smelt recruitment in the main basin of

Lake Huron, 1976-2010. Transaction of the American Fisheries Society, 143:784-

795.

17. Parker Stetter, S.L., Thomson, J.L.S., Rudstam, L.G., Parrish, D.L., Sullivan, P.J.,

2007. Importance and predictability of cannibalism in rainbow smelt. Trans. Am.

Fish. Soc. 136, 227–237.

18. Rupp, R.S. 1965. Shore-spawning and survival of eggs of the American smelt.

Transactions of the American Fisheries Society 94:160-168. 552

19. Scott, W.B., and E.J. Crossman. 1973. Freshwater fishes of Canada. Bulletin 184,

Fisheries Research Board of Canada, Ottawa.

20. Smith, P.W. 1979. The Fishes of Illinois. University of Illinois Press, Urbana,

Chicago, London.

553

CHAPTER 16

FAMILY PERCOPSIDAE

Introduction

Two species, the Trout-perch, Percopsis omiscomaycus, and the Sand Roller,

Percopsis transmontana, are the only two fishes deriving from the single extant genus,

Percopsis, that make up the members of Trout-perch family, Percopsidae. Both species are endemic to North America, and only the Trout-perch occurs in the Dakotas. The better known Trout-perch is distributed from Quebec through Hudson Bay to the Yukon in Alaska, south to Manitoba and the Upper Mississippi River basin to Kentucky, and east to the Hudson River. The less well known of the two species, the Sand Roller, has a much more limited distribution as it is endemic to the Columbia River drainage in

Washington, Oregon and Idaho where it is considered rare.

The family Percopsidae belongs to the relatively small order of ray-finned fishes

Percopsiformes. This order of fishes exhibits primitive, ancestral characteristics such as the presence of a small adipose fin, as well as more derived characteristics like fewer spines in the dorsal and anal fins. The common name of the family derived from both extant species sharing a combination of perch-like characteristics from the perch family

Percidae, such as a large head, teeth present, and a spinous first dorsal fin, and trout-like characteristics, such as an adipose fin that is typical of the trout family, Salmonidae. Both species of Trout-perch also have weakly ctenoid scales, and have rather large pectoral fins that overlap the pelvic fin when laid flat against the body. 554

Both species generally inhabit lotic environments, however they can also be successful in lentic environments. The Trout-perch often occurs at shallow and intermediate depths in areas with clear to slightly turbid water with slow to moderate current over sand or rocky substrates. They are also often associated with submerged structures such as woody debris and roots near undercut banks. Both species exhibit nocturnal behaviors, as they are most active when feeding during the night. Adults of both species feed on small aquatic insect larvae and occasionally other small minnows.

Trout-perch are known to spawn in the spring in shallow waters with sand, gravel or rocky substrate. Due to their small size, often not attaining more than 102mm (4in), they are a suitable prey item for larger predatory fishes.

555

Trout-perch, Percopsis omiscomaysus (Waldbaum, 1792)

Etymology and Synonyms: Percopsis = Greek, perke = perch + Greek, opsis = appearance; omiscomaycus = most likely an Algonkian Indian name that includes the root

‘trout’.9

Description: Body elongate, fusiform, cylindrical, but slightly compressed. Coloring translucent in appearance; dorsal and lateral sides olive-brown fading to a silvery-white belly; silvery stripe running along lateral line with 9 – 12 dark blotches; several indistinct dark spots present above the lateral line from head to tail. Fins lightly pigmented. Head large, slender and scaleless. Snout pointed, long, extending past mouth; upper lip groove not continuous. Mouth large and subterminal, slightly oblique to horizontal. Barbels absent. Teeth minute and villiform, numerous on upper and lower jaws. Dorsal fin positioned nearer tip of snout than base of caudal fin; large, slightly falcate with two soft spines and 9 –11 rays. Adipose fin present, small. Caudal peduncle long, tapered. Caudal fin forked. Anal fin with one soft spine, 5 – 8 rays. Pelvic fin originating posterior of dorsal fin insertion; one soft spine and 8 – 9 rays. Pectoral fin large, overlapping pelvic fin with 12 – 15 rays. Lateral line complete with 45 – 51 scales. Scales small and weakly ctenoid. Spawning adults and juveniles similar in appearance to adults.

Similar Species: No other species can be mistaken for Trout-perch in the Dakotas. A combination of ctenoid scales, adipose fin, and soft spines in the dorsal, anal, and pelvic fins make this fish unique. May be confused with juvenile Yellow Perch (Perca flavescens), with the adipose fin being an obvious distinguishing characteristic.

Distribution and Habitat: Distributed in North America from Quebec through Hudson

Bay to the Yukon in Alaska, south to Manitoba and the Upper Mississippi River basin to 556

Kentucky, and east to the Hudson River.6 Have only been collected in the Minnesota and

Big Sioux river drainages in northeastern South Dakota.1 Is said to have gained access to the Missouri River drainage by connection from the Minnesota River drainage.1 Range in

North Dakota includes upper and lower Sheyenne River, the Wild Rice River, the Souris

River, and the Red River drainage.8 Inhabits shallow and intermediate depths of clear to slightly turbid lakes and pools in large rivers and streams with slow to moderate current during the night. Prefers deeper water during the day. Found primarily over gravel and sand substrate. Known to inhabit deep glacial lakes.

Reproduction: Spawning occurs in spring from late April to June, or when water temperatures reach roughly 15.6 – 20°C (60 – 68°F).2 Spawning takes place over sand, gravel or rocky substrate a few inches below the surface. Sexual maturity is reached between ages 1 and 2, or 5.08 – 6.35cm (2 – 2.5 in.) TL.3 Males tend to mature and spawn at a younger age than females. Two to three males court a female over spawning grounds, most often at night. Females produce between 200 and 500 large eggs which descend to the bottom substrate and adhere. Eggs from Wisconsin were reported to be

1.87mm (0.07 in.) in diameter.2 Spawning area is abandoned and no parental care is given to the eggs or fry. Eggs hatch within roughly six days.

Age and Growth: Average adult length 76.2 – 101.6mm (3 – 4in.) TL. Larvae have been reported to be 5.3mm (0.21) TL.2 Wisconsin young of year reported at 27mm (1.06 in.)

TL in June, and 43 – 57mm (1.69 – 2.24 in.) in late September.2 Male lengths at age from

Lower Red Lake in Minnesota are: age 1, 50.8mm (2.0 in.); age 2, 88.2mm (3.47 in.); age

3, 103.5mm (4.07 in.).2,7 Adults capable of reaching 152.4 – 177.8mm (6 – 7 in.). 557

Specimens in South Dakota have not exceeded 76.2mm (3 in.).5 Can live to roughly 4 years.

Food and Feeding: Feeding takes place at night in shallower waters.8,9 Juveniles consume greater numbers of zooplankton than adults, consisting of ostricods, amphipods, rotifers, cladocerans and copepods.10 Adults consume mayfly and chironomid larvae, other insects and small minnows.4,8

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

4. Carlander, K.D. 1969. Handbook of freshwater fishery biology, volume 1. Iowa

State University Press, Ames.

5. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

6. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

7. Magnuson, J.J., and L.L. Smith. 1963. Some phases of the life history of the

troutperch, Percopsis omiscomaycus. Ecology 44(1):83-95.

8. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks. 558

9. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

10. Tomlinson, J.C., and D.J. Jude. 1977. Food of the trout-perch, Percopsis

omiscomaycus, in southeastern Lake Michigan. 20th Conf. Great Lakes Res., Int.

Assoc. Great Lakes Res., University of Michigan. Abstract.

559

CHAPTER 17

FAMILY GADIDAE

Introduction

The Burbot family, Gadidae, consists of 20 genera and 55 species worldwide, with roughly 16 species occurring in North America, and only one species, the Burbot,

Lota lota, occurring in the Dakotas. While the majority of the species within the family occupying cold, northern marine waters, the Burbot is the only species within the family

Gadidae to entirely inhabit cold, northern inland freshwaters. Cod are sometimes placed in the family Lotidae, which have 1-2 spine-less dorsal fins, whereas species in the family

Gadidae have 2-3 spine-less dorsal fins, with the exception of the Burbot, which has 2.

Cod have a rather distinct appearance from other fishes, especially in the Dakotas, by their elongate, cylindrical bodies that taper posteriorly on the body to create a slightly laterally compressed appearance. Cod can also have 1-2 anal fins, with the Burbot displaying a single, elongate anal fin giving it an eel-like appearance. The majority of species with the Cod family can also be recognized and distinguished from other families of fishes with elongate body shapes, like the eel family, Anguillidae, by the single barbel present on the chin. Cod also have extremely small and embedded cycloid scales, which give them a scale-less look, much like catfish.

As stated previous, the majority of Cod species spend their entire lives inhabiting marine waters, with very few rarely entering freshwaters for short periods of time. The

Burbot is the one lone species in the family to undergo its entire life cycle in freshwater.

Burbot prefer the deep and cold waters of large lakes, reservoirs, and rivers, which 560 explains it distribution throughout the Missouri River and Red River of the North in the

Dakotas. They are often found hiding within dark crevices or under cover during the day, and are most active at night when they come out to feed. As larvae and juveniles, Burbot forage on phytoplankton, zooplankton, and small aquatic insects before they undergo an ontogenetic diet shift to piscivory as adults. Spawning of Burbot takes place in the winter and early spring right under the ice, generally at night. Individuals may not spawn every year, but when they do, spawning takes place in groups that are often described as “ball- like”.

In the more northern latitudes, Cod are an important group of commercial and highly sought after fishes, and serve as an important and delightful food source for many anglers. In the Dakotas, Burbot are less popular for sportfishing and are sometimes even considered a rough fish. Although their distribution is rather widespread throughout the northern waters of North America, Europe and Asia, many populations of Burbot are declining due to the construction of impoundments and pollution.

561

Burbot, Lota lota (Linnaeus, 1758)

Etymology and Synonyms: Lota = French for “la Lotte”, meaning “cod” or “codfish”; lota = repeats the genus translation emphasizing “cod”. The common name, “Burbot” stems from the Latin word “barba”, meaning “beard”, referring to the single barbel or whicker that stems from the chin.

Description: Body elongate, anteriorly cylindrical, laterally compressed posteriorly.

Dorsal and lateral coloration variable, and may be mottled or rather solid mixtures of dark brown, black, tan, or brownish-yellow; ventrally light yellow, cream or white. Head small; moderately dorsoventrally compressed. Nostrils small and tube-like; located anteriorly on head. Eye small, positioned laterally on head. Single barbell present, located on ventral side of chin. Mouth large, wide; maxilla slightly extends over mandible. Teeth small, villiform, arranged in bands on both jaws. Dorsal fin distinctly divided into two sections; anterior dorsal fin short with 8-16 rays; posterior dorsal fin short, extending nearly half the length of the body 60-80 rays. Caudal fin distinctly separated brom dorsal and anal fins; rounded distal end. Anal fin nearly as long as posterior dorsal fin with roughly 53-65 rays; insertion posterior to posterior dorsal fin insertion. Pelvic fins narrow with 5-8 rays; second fin ray elongated; insertions anterior to insertion of pectoral fins.

Pectoral fins with 17-21 rays and rounded distal ends. Lateral line complete with numerous small, embedded cycloid scales in series; appears scaleless.

Similar Species: The American Eel is the only species in the Dakotas that slightly resembles the Burbot due to its larger, elongate, and slender body shape. American Eel can easily be distinguished from Burbot by the lack of a barbel under the chin, and having continuous, fused dorsal, caudal, and anal fins. Juvenile Burbot may also could be 562 mistaken for smaller juvenile bullheads or catfishes, but again can be easily identified by the presence of a single barbel on the chin.

Distribution and Habitat: The Burbot is the only representative of the cod family,

Gadidae, that completes its entire lifecycle in freshwater.12 However, in some areas it is known that Burbot have been identified in estuaries and brackish lagoons.8 Burbot are a holarctic species, with a circumpolar native range extending across northern Europe,

Asia, and North America across Canada, Alaska and the northern United States.8,10

Within North America, it ranges as far south as Missouri, and in the Dakotas, it primarily occurs within the Red River of the North and the Missouri River basins.1,8 Burbot prefer deep, cold, rather dark waters of lakes, reservoirs, large rivers and streams with a variety of substrates. They often hide within dark crevices under boulders or any other available cover. Juveniles often inhabit areas along shorelines, in weed beds, or under cover in little to swift current.7

Reproduction: Burbot spawn in lakes, reservoirs, rivers, and streams, often in the winter or early spring under the ice. The spawning season generally lasts 2-3 weeks. Colder water temperatures are preferred, and the time of day spawning occurs may be dependent on the region, but spawning predominately occurs at night.8,9 Spawning has been described as a large “ball” with one or two females in the center releasing eggs who are surrounded by many males releasing sperm.8 Not all adults spawn every year; “skipping” spawning is particularly common in northern populations.8 Fecundity varies regionally, but is often large (6,300-3,477,699 eggs) and is positively correlated with the age and size of the female.8,5 Eggs round, roughly 1 mm (0.04 in) in diameter, semi-buoyant to 563 buoyant, and non-adhesive. Eggs are known to develop faster at higher temperatures.8

Larval Burbot hatch near the substrate before becoming pelagic.7

Age and Growth: Mean length of larval Burbot captured in North Dakota during spring was 4.9 mm (0.19 in) TL in April and 80.9 mm (3.19 in) in July.2 Juveniles may reach

110-120 mm (4.33-4.72 in) TL by late fall.2,8,11,12 Growth in the first year is rapid and varies with length of growing season and food resources. Average length of age-1 Burbot from North Dakota was 152.6 mm (6.00 in) TL.2 Burbot typically grow to 1 m (39.37 in)

TL and weigh roughly 8 kg (17.637 lbs.); however the median size is 300-600 mm

(11.81-23.62 in) TL and 1-3kg (2.20-6.61 lbs.).8 Burbot populations from rivers and reservoirs generally have lower relative weight (Wr) values across length categories than populations within lakes.3,7 Record-size Burbot in Siberia grow to 25-30 kg (55.11-66.14 lbs.) and ages of 15-20 years.8 Maximum age in a northern population is 22 years.8

Food and Feeding: When larvae are big enough to drift through the limnetic zone, they are known to select larger zooplankton such as rotifers, copepods, and cladocerans.2,4

Juveniles and smaller individuals continue to forage on zooplankton, but will also begin to consume crayfish, macroinvertebrates including Odonata, Amphipoda, and Plecoptera, and small benthic fishes.2,6 Burbot undergo and ontogenetic diet shift and become increasingly piscivorous and a top level predator as they grow in size and reach adulthood.6

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor. 564

2. Fisher, S.J. 2000. Early life history observation of Burbot utilizing two Missouri

River backwaters. Burbot biology, ecology, and management. American Fisheries

Society, Fisheries Management Section, Publication, Bethesda, Maryland, 96-0.

3. Fisher, S.J., D.W. Willis, and K.L Pope. 1996. An assessment of Burbot (Lota

lota) weight-length data from North American populations. Canadian Journal of

Zoology 74:570-575.

4. Ghan, D., and W.G. Sprules. 1993. Diet, prey selection, and growth of larval and

juvenile Burbot Lota lota (L). Journal of Fish Biology 42: 47-64.

5. Hesse, L. W. 1993. The status of Nebraska fishes in the Missouri River. 2. Burbot

(Gadidae: Lota lota). Transactions of the Nebraska Academy of Sciences and

Affiliated Societies. Paper 120.

6. Jacobs, G.R., C.P. Madenjian, D.B. Bunnell, and J.D. Holuszko. 2010. Diet of

Lake Trout and Burbot in Northern Lake Michigan during spring: evidence of

ecological interaction. Journal of Great Lakes Research 36:312-317.

7. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

8. McPhail, J.D. and V.L. Paragamian. 2000. Burbot biology and life history.

Burbot: biology, ecology, and management. American Fisheries Society, Fisheries

Management Section, 1:11-23.

9. Roach, S.M. and M.J. Evenson. 1993. A geometric approach to estimating and

predicting fecundity of Tanana River burbot. Alaska Department of Fish and

Game, Fisheries Data Series No. 93-38, Juneau. 565

10. Ryder, R.A., and J. Pesendorf. 1992. Food, growth, habitat, and community

interactions of young-of-the-year burbot, Lota lota L., in a Precambrian Shield

lake. Hydrobiologia, 243/244: 211-227.

11. Sandlund, O.T., L. Klyve, and T.F. Naesje. 1985. Growth, habitat and food of

Burbot, Lota lota, in Lake Mjoesa. Fauna Blindern 38: 37-43 (in Norwegian with

English summary).

12. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fisheries

Research Board of Canada, Bulletin 184. 966p.

566

CHAPTER 18

FAMILY FUNDULIDAE

Introduction

The Topminnow and Killifish family, Fundulidae, is made of four genera and roughly 40 species that are native to North and Central America from Canada to Cuba and the Bahamas. Three species of Fundulids occur in the Dakotas: the Banded Killifish,

Fundulus diaphanous, the Northern Plains Killifish, Fundulus kansae, and the Plains

Topminnow, Fundulus sciadicus. These three species, along with many others in the family, are rather small in size and display distinct features that help separate themselves from species in the minnow and carp family, Cyprinidae. Fundulids have elongate, slightly laterally compressed bodies that are dorsoventrally flattened on the head and across the anterior end of the dorsal side. The mouths of Fundulids are also small and upturned, or superior with the lower jaw protruding past the upper jaw. Fundulids origionate from the order Cyprinodontiformes, which is Greek for “toothed carp”, referencing the many small, unicuspid teeth placed in one or more rows on both the upper and lower jaws. The dorsal fin of Fundulids lacks any spines and is placed far posterior on the body above the anal fin. Fundulids can also easily be distinguished from Cyprinids by the shape of the caudal fin, which has a rounded distal end, versus forked like minnows and carps.

Fundulids within the Dakotas occur in a variety of habitat types including small sized rivers and streams, ponds, sloughs, and lakes in relatively shallow areas with low velocity. Some species prefer clear water over silt and sand substrates with dense aquatic 567 vegetation nearby, like the Plains Topminnow, whereas other species such as the

Northern Plains Killifish and the Banded Killifish are more tolerant of turbidity and are often found over sand and gravel substrates. Together, the body shape and the position of the mouth have assisted Fundulids to adapt a unique foraging style, which includes skimming just beneath the surface of the water to feed on a variety of small aquatic invertebrates. Fundulids often display nuptial coloration during the spawning season, where males are often more colorful than females. Spawning habits are known to differ by species, but in general males become rather defensive of their spawning territories or other females, and no nest or parental care is given to the eggs are larvae pre- or post- hatching.

Some species within the Great Plains region are experiencing decreases in their historic range and population declines due to the loss and alteration of their habitat, as well as the range expansion of invasive species. Although their rather small size does not make them suitable for sportfishing, Fundulids often serve as forage species for piscivorous game fishes such as bass and pikes. They are also known to adapt easily and serve as amusing aquarium species.

568

Banded Killifish, Fundulus diaphanus (Lesueur, 1817)

Etymology and Synonyms: Fundulus = Latin for fundus, meaning “bottom”; a peculiar name for a species in the top minnow family, Fundulidae; diaphanous = Greek for transparent. The common name Banded Killifish refers to distinct, dark vertical bands along their lateral sides.

Description: Banded Killifish have a fusiform, elongate body that is laterally compressed. Dorsally olive green; laterally olive green to silver displaying 12-20 greenish-brown vertical bands; throat greenish-yellow; fins greenish-yellow to clear; ventrally silvery white. Head flattened; snout bluntly pointed. Mouth small and superior; lower jaw protruding further than upper jaw; continuous groove between snout and upper lip; teeth small and sharp, present in small rows along both jaws. Eye larger; placed medial on the lateral side of head. Dorsal fin base anterior to anal fin base with (10-15) rays; spines absent. Caudal fin homocercal; slightly rounded to convex. Anal fin with 10-

12 rays. Pelvic fins with 6 rays. Pectoral fins rounded. Lateral line absent; 39-49 larger cycloid scales in lateral series. Peritoneum silver. Sexual dimorphism present; females display darker bands along lateral sides than males.2 Spawning males develop darker vertical bands on lateral sides during spawning season.1,3

Similar Species: Northern Plains Killifish also have dark vertical bands along the lateral sides, however they have smaller scales with roughly 50-67 scales in the lateral series and a black peritoneum.3 The Central Mudminnow and the Western Mosquitofish also may be mistaken for Banded Killifish, except they both display a dark spot below the eye.

Distribution and Habitat: Banded Killifish are widely distributed throughout eastern

North America from Newfoundland to South Carolina, and west to the Great Lakes 569 drainages, all the way to the eastern side of the Dakotas. The species is native to the

Great Lakes and Mississippi River drainages, including portions of northern Iowa and northeastern Nebraska.3 In the Dakotas area, it was accidentally introduced into Lake

Andes, South Dakota while stocking Largemouth Bass.1 It has been known to inhabit the

Big Sioux river drainage from which it most likely gained access through the Minnesota and Des Moines river basins.1 It has also been reported in the Sheyenne River, Turtle

River, and headwaters of the Red River tributaries in North Dakota.4,5,6 Banded Killifish prefer quiet and shallow waters of sloughs, marshes, ponds and lakes, as well as low gradient brooks and streams with gravel or sand substrate and abundant vegetation.7,8,9

Juveniles school within aquatic vegetation, while adults are frequently seen in schools venturing out during the day into open water just inches below the surface. It is at this point where they expose themselves to predation by other piscivorous species like

Largemouth Bass, Smallmouth Bass, and Northern Pike.3,10 The species is tolerant to a range of salinities, low oxygen levels, and warmer water temperatures, making it adaptable to the plains region.

Reproduction: Sexual maturity is known to occur between 1-2 years of age, and is more likely known to be a function of growth than age.10,11 The spawning period is driven by water temperature, with preferred temperatures ranging from 21°C to 23°C (69.8°F to

73.4°F), and continues for roughly three weeks.10,11 Spawning behavior begins when females suspend a small cluster of eggs from the genital papilla via adhesive threads.

Males with bright breeding colors then pursue and court the females into a vegetated or grassy area within their territory in which they defend.4,13 The cluster of eggs is then further suspended and fertilized by the male. The eggs detach from the female and adhere 570 to the vegetation where they develop and hatch.4 Fecundity has been known to vary greatly depending on location. In two Canadian lakes, fecundity was found to be between

88 and 128 eggs, while another two lakes in Quebec Canada found fecundity to range between 226 eggs in an 82 mm (3.23 in) female, and 426 eggs in a 101 mm (3.98 in) female.10,11,12

Age and Growth: Adults average 6-8 cm (2.36-3.15 in) TL, but are capable of reaching

10-13 cm (3.94-5.12 in) TL.3 Banded Killifish are known to live up to 4 years old, with most adults surviving until age 3. Length-at-age form Lake Erie reported as: age-1, 52-67 mm (2.05-2.64 in) TL; age-2, 63-79 mm (2.48-3.11 in) TL; age-3, 93 mm (3.66 in) TL.10

Food and Feeding: Banded Killifish of all ages feed throughout the water column primarily on Cladocerans, benthic macroinvertebrates, and aquatic insect larvae.4,10,13

Juveniles feed primarily on Cladocerans and Chironomids. Adults broaden their diet by consuming Ostracods, Copepods, Odonata and Emhemeroptera nymphs, and some mollusks and Amphipods, but still continue to consume large volumes of Cladocerans.4,10

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Forbes, S.A., and R.E. Richardson. 1920. The fishes of Illinois. Illinois State

Journal Company, State Printers, Springfield.

3. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence. 571

4. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

5. Woolman, A.J. 1896. Report on ichthyological investigations in western

Minnesota and eastern North Dakota. Rept. U.S. Fish Comm. 1893:343-373.

6. Copes, F.A. 1965. Fishes of the Red River tributaries of North Dakota. M.S.

Thesis, University of North Dakota. 65 p.

7. Eddy, S., and J.C. Underhill. 1974. Northern fishes with special reference to the

upper Mississippi Valley. Third edition. University of Minnesota Press,

Minneapolis. 414 p.

8. Trautman, M.B. 1981. The fishes of Ohio, revised edition. Ohio State University

Press, Columbus.

9. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

10. Phillips, E.C., Y. Ewert, and P.A. Speares. 2007. Fecundity, age and growth, and

diet of Fundulus diaphanous (Banded Killifish) in Presque Isle Bay, Lake Erie.

Northeastern naturalist, 14(2):269-278.

11. Fournier, P., and E. Magnin. 1975. Reproduction du petit barré de l’est Fundulus

diaphanous diaphanous (Le Sueur). Naturaliste Canada 102:181-188.

12. Fritz, E.S., and E.T. Garside. 1975. Comparison of age composition, growth, and

fecundity between two populations each of Fundulus heteroclitus and F.

diaphanous (Pisces: Cyprinodontidae). Canadian Journal of Zoology 53:300-311.

13. Stewart, K.W. and Watkinson, D.A. 2004. The freshwater fishes of Manitoba.

University of Manitoba Press, Winnipeg.

572

Plains Topminnow, Fundulus sciadicus (Cope, 1865)

Etymology and Synonyms: Fundulus = Latin for fundus, meaning “bottom”; sciadicus

= “shady”, referring to the preferred habitat of the species.

Description: Plains Topminnow have a moderately robust and stout body anteriorly, and are laterally compressed posteriorly. Dorsally dark olive green with prominent gold dorsal stripe anterior of dorsal fin insertion; laterally olive green to silver; ventrally silver; fins faint yellow-orange to clear and lacking vertical bands on lateral sides; top of head with small silvery-blue specks present. Head large, broad, and dorsoventrally flattened with scales present. Snout blunt. Eye moderately large with gold iris; placed laterally on head. Mouth small, superior; lower jaw slightly protrudes past maxillary.

Teeth small, present on both jaws. Dorsal fin positioned posteriorly on body with slightly rounded distal end; 9-11 rays, spines absent. Adipose fin absent. Caudal peduncle short and thick. Caudal fin homocercal with slightly rounded distal end. Anal fin insertion slightly anterior to insertion on dorsal fin; elongate with round distal end and 12-15 rays.

Pelvic fin small, placed abdominally. Pectoral fins with rounded distal end. Lateral line absent; 33-37 large cycloid scales in lateral series. Spawning males with nuptial coloration; dorsally dark blue-green with bright red-orange coloring in fins.

Similar Species: Resembles the Central Mudminnow. Central Mudminnow display an anal fin with only 7-9 rays, 34-37 larger cycloid scales in the lateral series, and a prominent dark vertical bar at the base of the caudal fin. Banded Killifish and Northern

Plains Killifish display dark vertical bands on lateral sides.

Distribution and Habitat: Endemic to the Great Plains region in the central United

States, particularly in two isolated populations. The largest expands from eastern 573

Nebraska, north to the northwestern corner of Iowa, the southern range of Minnesota and

South Dakota, and west to northeastern Colorado and southeastern Wyoming.5 The second smaller population extends from central Missouri, to eastern Kansas and the northeast corner of Oklahoma.5 In South Dakota, the Plains Topminnow is found in the tributaries of the Cheyenne, Keya Paha, Little White, Big Sioux, Niobrara, Vermillion, and James River basins.6 Absent from North Dakota. The species has experienced as much as a 70% decrease in its historic range, vastly due to the loss and alteration of habitat as well as the expansion of the invasive Western Mosquitofish.5,7,10 Considered a habitat specialist due to specific habitat requirements. Plains Topminnow are found in clear headwater streams with sand, silt, or mud substrate with dense aquatic vegetation.11,15 They also prefer quiet spring-fed shallowpools, sloughs, and backwaters of slightly larger streams. Optimum water temperature 18.3-23.8°C (65-75°F), with a mean critical thermal maximum of 37°C (98.6°F).2,13 Minimum home range is 1,300 lineal meters (4,265 ft.).12 Tolerant of low dissolved oxygen concentrations, and elongated periods with high water temperatures in isolated pools.13

Reproduction: Spawning is temperature dependent and varies with location, however it generally occurs late March through early August for approximately 60 days when water temperatures reach 18-24°C (64.4-75.2°F).3 Sexual maturity is reached at ages 1-2, leaving both males and females sexually mature for roughly 3 years.3,11,14 During spawning, males become defensive of other males and display nipping behavior.3 Males attract and court females to spawning grounds by enlarging their gular region.3 Spawning activity consists of the “wiggle” or “tilt” behaviors, in which males rub and press their bodies vigorously on the females against vegetation or the substrate to release and 574 fertilize eggs.1,3 No nest is built and no parental care is given. Eggs are broadcast and adhere to detritus and vegetation. Eggs are yellow-orange in color and roughly 0.1-1.8 mm (0.004-0.071 in) in diameter.3,4 All eggs released at one time.8 Fecundity rather low; average 50-90 eggs per female.2 Hatching occurs within 13-14 days at 21-23°C (69.8-

73.4°F).3,4

Age and Growth: Newly hatched larvae are typically 6.2-7.7 mm (0.24-0.30 in) TL.3

Mean lengths-at-age from Nebraska are reported as: age-0, 23 mm (0.91 in) TL; age-1,

47 mm (1.85 in) TL; age-2, 53 mm (2.09 in) TL; age-3, 62 mm (2.44 in) TL.14 Average size 32-64 mm (1.26-2.52 in) TL.8,9 Capable of reaching 75 mm (2.95 in) TL.8 Females slightly larger in length and weight than males of same year class.14 Longevity is 4 years.

Food and Feeding: Information on feeding habits limited. Feeding primarily takes place at the surface. Preferred feeding habitat consists of heavily vegetated backwater habitat.16

Adults mainly consume a variety of small aquatic invertebrates including adult and larval forms of Diptera, Hemiptera, and occasionally small crustaceans and snails.8,9,14

Individuals in lentic environments have shown to demonstrate a generalist feeding strategy, while lotic individuals favor Gastropods over Decapods and Ephemeropterans.16

In Missouri, diets utilized a greater amount of benthic organisms, indicating that diets may vary due to location, season, and trophic interactions.16,17

Literature Cited:

1. Baugh, T.M. 1981a. In search of Plains Topminnow. Freshwater and Marine

Aquarium 4(9):39-41.

2. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL. 575

3. Kaufmann, S.A., and J.D. Lynch. 1991. Courtship, eggs, and development of the

Plains Topminnow in Nebraska (Actinopterygii: Fundulidae). Prairie Naturalist

23(1):41-45.

4. Kinney, T.A., and J.D. Lynch. 1991. The fecundity and reproductive season of

Fundulus sciadicus in Nebraska (Actinopterygii: Fundulidae). Transactions of the

Nebraska Academy of Sciences 18:101-104.

5. Pasbrig, C.A. 2010. Reductions in range-wide distribution of Plains Topminnow,

Fundulus sciaicus, and production of a broodstock pond. M.S. thesis, University

of Nebraska, Kearney.

6. Pasbrig, C.A. 2012. Plains Topminnow: the “minnow” that isn’t a minnow. South

Dakota Conservation Digest 79(5):2-3.

7. Pasbrig, C.A., K.D. Koupal, S. Schainost, and W.W. Hoback. 2012. Changes in

range-wide distribution of Plains Topminnow Fundulus sciadicus. Endangered

Species Research 16:235-247.

8. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

9. Rahel, F.J., and L.A. Thel. 2004. Plains Topminnow (Fundulus sciadicus): A

technical conservation assessment. USDA Forest Service, Rocky Mountain

Region, Golden Colorado.

[http://www.fs.fed.us.r2/projects/scp/assessment/plainstopminnow.pdf.]

10. Schumann DA, Hoback WW, Koupal KD. 2015. Complex interactions between

native and invasive species: investigating the differential displacement of two

topminnows native to Nebraska. Aquatic Invasions 10:339-346. 576

11. Schumann, D.A., C.A. Pasbrig, K.D. Koupal, and W.W. Hoback. 2012. Culture of

Plains Topminnow in a pond constructed for species conservation. North

American Journal of Aquaculture 74(3):360-364.

12. Schumann, D.A., K.D. Koupal, W.W. Hoback, C.W. Schoenebeck, and S.

Schainost. 2014. Large-scale dispersal patterns and habitat use of Plains

Topminnow Fundulus sciadicus: implications for species conservation. Journal of

Freshwater Ecology 30:311-322.

13. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

14. Stribley, J.A., and R.H. Stasiak. 1982. Age, growth and food habits of the Plains

Topminnow, Fundulus sciadicus Cope, in Keith County, Nebraska. Proceedings

of the Nebraska Academy of Sciences and Affiliated Societies 92:17-18.

15. Thiessen JD. 2016. Conservation of Plains Topminnow, Fundulus sciadicus,

reestablishment success and limiting factors of persistence of reintroduced

populations in Nebraska. M.S. Thesis, University of Nebraska at Kearney.

16. Thiessen, J., K. D. Koupal, C. W. Schoenebeck, and J. J. Shaffer. 2018. Food

habits of imperiled Plains Topminnow and diet overlap with invasive Western

Mosquitofish in the central Great Plains. Transactions of the Nebraska Academy

of Sciences and Affiliated Societies 38:1-9.

17. Thompson, G.T. 2014. Ecology of a declining Great Plains fish, Fundulus

sciadicus, in the Missouri Ozarks. M.S. thesis, Missouri University of Science and

Technology, Rolla. 577

Northern Plains Killfish, Fundulus kansae (Garman, 1895)

Etymology and Synonyms: Fundulus = Latin for fundus, meaning “bottom”, referring to the associated habitat of the species when feeding; kansae = referring to the locality of the species.

Description: Northern Plains Killifish have a fusiform, elongate body that is laterally compressed. Dorsally olive to tan; laterally light olive, silver, or cream with 12-28 dark gray vertical bars; ventrally cream to white; fins without markings, clear to light gray.

Head large, flattened dorsally with large scales present. Snout elongate. Eye large, placed dorso-laterally on head. Mouth small, superior; lower jaw protrudes past maxillary. Lip on lower jaw large, fleshy. Teeth small, fine, villiform. Dorsal fin positioned posteriorly on body with rounded distal end; 13-15 rays. Adipose fin absent. Caudal peduncle thick, elongate. Caudal fin homocercal with nearly straight distal end. Anal fin insertion directly beneath dorsal fin insertion; large with round distal end and 13-14 rays. Pelvic fin small, placed abdominally. Pectoral fins with rounded distal end. Lateral line absent; 47-67 small cycloid scales in lateral line series. Peritoneum black. Intestine long. Sexual dimorphism present; males with fewer and wider vertical bars on lateral sides than females. Spawning males develop darker vertical bars on the lateral sides, as well as bright red-orange coloring in the caudal, anal, pelvic, and pectoral fins.

Similar Species: Closely resembles the Banded Killifish. Banded Killifish also display dark vertical bands on the lateral sides, however they have larger cycloid scales with roughly 39-49 scales in the lateral series, and a silver peritoneum. Central Mudminnow display an anal fin with only 7-9 rays, and 34-37 larger cycloid scales in the lateral series.

Plains Topminnow lack vertical bars on lateral sides. 578

Distribution and Habitat: Native to the Great Plains region in North America in the

Arkansas and Missouri River drainages, from Montana in the west, east to Missouri, and south to Texas. Occurs in South Dakota throughout the Cheyenne River drainage and its major tributaries.7 Absent from reservoirs within the Cheyenne River drainage.7 Due to habitat alterations, the species is declining in parts of its range creating a patchy distribution.3 Often found schooling within medium to small sized rivers and streams, as well as small ponds, shoals, and backwaters with low flow in lower elevations.11,12 Prefer shallow water over sand or gravel substrate.3 Avoids areas deeper than 15 cm (5.90 in) in depth.9 Tolerant to high salinity, alkalinity, and intermittency.1,10 Avoids areas with high levels of siltation.6 Northern Plains Killifish have also been observed burying themselves within sand substrate leaving only their eyes and mouth visible.8 This behavior may possibly be performed to escape predators, create camouflage when searching for food, or to seek cooler temperatures.8

Reproduction: Both sexes reach sexual maturity at age-1, with the majority reaching sexual maturity at age-2.9 Females begin developing eggs in February.9 Spawning takes place in summer months, typically April-August, or when water temperatures near

25.5°C (78°F). Spawning grounds consist of shallow waters less than 10 cm (3.94 in) deep with sand or gravel substrate and low current.9 Males known to defend small territories from other males and also compete for females prior to spawning. A single male actively courts one female to the spawning site where their bodies vibrate until eggs are released and fertilized. No nest is prepared and no parental care is given; adults retreat back to preferred habitat soon after spawning concludes. Eggs roughly 2.065 mm (0.08 579 in) in diameter.9 Fecundity is low and increases with size of female; 10-100 eggs per female.2,9 In aquariums, hatching occurred within 2-3 weeks.9

Age and Growth: Newly hatched larvae average 9.53 mm (0.38 in) TL.2 Roughly 35 mm (1.38 in) TL by the end of their first summer, and 50 mm (1.97 in) TL at the end of their second summer.9 Capable of reaching 100 mm (3.94 in) TL.4 Longevity is 2 years.

Food and Feeding: Primarily carnivorous. Feeds by sight throughout the water column during the day. Also known to be a benthic feeder and digs its snout into the substrate taking a mouthful, retaining any organisms.9 Feeding generally does not take place at night.5 Adults consume a variety of prey items including insects and aquatic invertebrates and larvae such as Ephemeroptera and Chironomidae.4,9 Also occasionally consume plant material and diatoms.

Literature Cited:

1. Baxter, G.T., and M.D. Stone. 1995. Fishes of Wyoming. Wyoming Game and

Fish Department, Cheyenne.

2. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

3. Brown, K.L. 1986. Population demographic and genetic structure of Plains

Killifish from the Kansas and Arkansas River Basins in Kansas. Transactions of

the American Fisheries Society 115:568-576.

4. Eberle, M.E. 2009. Type locality and conservation status of the Northern Plains

Killifish (Fundulus kansae: Fundulidae) in Kansas. Transactions of the Kansas

Academy of Science 112:87-97. 580

5. Echelle, A.A., M.M. Stevenson, A.F. Echelle, and L.G. Hill. 1971. Diurnal

periodicity of activities in the Plains Killifish, Fundulus zebrinus kansae.

Proceedings of the Oklahoma Academy of Science 51:3-7.

6. Gido, K.B., C.S. Guy, T.R. Strakosh, R.J. Bernot, K.J. Hase, and M.A. Shaw.

2002. Long-term changes in the fish assemblages of the Big Blue River Basin 40

years after the construction of Turtle Creek Reservoir. Transactions of the Kansas

Academy of Science 105:193-208.

7. Hoagstrom, C.W., C-A. Hayer, and C.R. Berry. 2009. Criteria for determining

native distributions of biota: the case of the Northern Plains Killifish in the

Cheyenne River drainage, North America. Aquatic Conservation: marine and

Freshwater Ecosystems 19:88-95.

8. Minckley, C.O., and H.E. Klaassen. 1969a. Burying behavior of the Plains

Killifish, Fundulus kansae. Copeia 1969:200-201.

9. Minckley, C.O., and H.E. Klaassen. 1969b. Life History of the Plains Killifish,

Fundulus kansae (Garman) in the Smokey Hill River, Kansas. Transactions of the

American Fisheries Society 98:460-465.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

11. Quist, M.C., W.A. Hubert, and F.J. Rahel. 2004. Elevation and stream-size

thresholds affect distributions of native and exotic warmwater fish in Wyoming.

Journal of Freshwater Ecology 19:227-236. 581

12. Senecal, A.C. 2009. Fish assemblage structure and flow regime of the Powder

River, Wyoming: an assessment of the potential effects of flow augmentation

related to energy development. M.S. thesis, University of Wyoming, Laramie.

582

CHAPTER 19

FAMILY GASTEROSTEIDAE

Introduction

The Stickleback family, Gasterosteidae, is made of five genera and roughly 16 species. Sticklebacks have a Holarctic distribution and are spread across North America,

Asia and Europe, with roughly six species occurring in the United States. Most species in the family are marine and prefer coastal and brackish waters, however, the only species in the genus Culaea, the Brook Stickleback, Culaea inconstans, is the only freshwater member in the family, and the only species of Stickleback that occurs in the Dakotas.

Sticklebacks are a group of small, scaleless fishes that have an ovate and laterally compressed body shape. The dorsal spines, each followed by a small, trailing membrane, easily distinguish themselves and give the family a unique appearance among other groups of fishes, especially across the Great Plains region. Sticklebacks also have slightly elongate and narrow caudal peduncles, a fan-shaped caudal fin, and a rayed dorsal fin that sits far posterior on the body, behind the dorsal spines. Although Sticklebacks are scaleless, some species, including the Brook Stickleback, are suited with weakly developed dermal or body plates along the body.

Members of the stickleback family have rather intricate and unique life history traits that are species specific. For example, the Three-spined Stickleback, Gasterosteus aculeatus, which does not occur within the Dakotas but is spread throughout the Great lakes region as well as the Pacific and Atlantic coasts, has been the subject of many scientific studies due to their captivating social, spawning, and anti-predator behaviors. 583

Many species of Stickleback, including the Brook Stickleback, display interesting spawning and nest building behavior. All male sticklebacks construct a nest that often times are secured to submerged aquatic vegetation and are held together by secretions produced from the males kidneys. Nests are hollow on the inside, and have one small hole for the males to lure females into. Once spawning commences inside the hollow nests, females of most species create a hole in the back of the nest to exit, which the male later patches. Males also display parental care by guarding and tending to the eggs and larvae pre- and post- hatching.

Sticklebacks often times serve as suitable prey for various piscivorous sport fishes such as sunfish, pikes and salmon. They are also consumed by a wide array of avian predators. Their unique appearance combined with their intricate life history traits makes sticklebacks an amusing group of aquarium fishes.

584

Brook Stickleback, Culaea inconstans (Kirtland, 1840)

Etymology and Synonyms: Culaea originates from the Greek name “Eucalia” meaning good nest, referring to the Brook Sticklebacks intricate nest structure; inconstans stands for variable. Brook Stickleback are the only freshwater member in the stickleback

(Gasterosteidae) family, as well as the only species in its genus (Culaea). Other common names for this species are Five-spined Stickleback and Common Stickleback.

Description: Unique appearance among plains fishes. Body ovate and laterally compressed. Head moderately pointed. Mouth small and oblique; teeth sharp and located in narrow band on jaw only. Eye large. Scales absent; replaced by a single, lateral row of bony plates, or dermal bones.1,6,8 Most noted for their dorsal spines with small, trailing membranes; typically five spines, but can vary from 2-7 and 9-11 rays.8 Lateral line complete. Adults usually display a dark tessellated pattern on the dorso- and lateral sides; often have a yellow, to olive green color background with a lighter lateral surface than the ventral.8 Brook Stickleback are able to rapidly change color to camouflage with the background conditions or to show aggressiveness.8 Caudal peduncle narrow. Anal fin with one spine and 10-11 rays. Pelvic fin with one spine and no rays. Pectoral fin absent of spines but includes 9-11 rays. There is no sexual dimorphism outside of breeding season. Breeding males display much darker green to jet black coloring on their dorsal side; they also acquire a black vertical band of enlarged melanophores within their bright yellow iris of the eye for the duration of the breeding season and especially during territorial encounters.8

Similar Species: The Brook Stickleback is unlikely to be easily confused or misidentified with any other species in North Dakota and South Dakota. 585

Distribution and Habitat: Range within the United States spans from eastern Montana to New York and northern Maine, and south to the Ohio River system in Pennsylvania,

Ohio and Indiana, the Illinois River system in Illinois, and the Missouri River system in

Kansas.1,6 Has been reported in all drainages of North Dakota.4 Fairly abundant in the eastern part of South Dakota, often occurring in cool and clear lakes and streams.4,5

Known to be present west of the Missouri River in South Dakota, although not as abundant due to diminishing habitat needs.6 The Brook Stickleback is one of the most cold adapted freshwater fishes and prefers to inhabit cool and clear vegetated waters, often headwater streams, ponds, and lakes.8 They are known to be fairly tolerate to salinity, however do not do well with high and prolonged levels of turbidity and water temperature.3,4,8

Reproduction: Spawning can be prolonged and occur during the summer months, or when water temperatures reach roughly 46-66°F.2 Brook Stickleback are often recognized for their intricate and amusing spawning habits. Sexual maturity is most likely reached during the second growing season.3 Males construct and defend a spherical and hollow nest the size of a walnut with one opening. It has been noted that Brook Stickleback are vigorous fishes, and will act defensive towards larger fish.5 Nests are made from filamentous algae and other aquatic vegetation, and are often attached to twigs or rooted aquatic plants from secretions produced by the male’s kidney and other organs. Males lure females into the back of the nest, and rub against them to release her eggs. Females are known to release 100-300 eggs at a time. Once the eggs are released, males are known to block the entryway to prevent eggs from escaping. The female then creates an opening in the back of the nest to escape. Once the female leaves, the male fixes the 586 opening and continues to lure other females into the nest. Eggs hatch within 8-9 days and fry are given parental care by the male.2 Fry are roughly one-quarter inch in length. If fry try to leave the nest too soon, males will herd them back into the nest. When the fry become too much to handle, the male will then abandon the nest, or feed on the fry.2

Age and Growth: The length of Brook Stickleback does not often exceed 90mm TL.7

Fry hatch at roughly one-quarter inch, and grow to roughly 25-51mm their first year, and

51-76mm during their second year.2,3,4 Individuals have been known to live for roughly 3 years.2

Food and Feeding: Brook Stickleback are primarily carnivorous, feeding mainly on small aquatic insects, insect larvae, fish eggs and fry, worms, and algae. They have also been known to consume small crustaceans and snails.4,5

Literature Cited:

1. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

2. Brown, C.J.D. 1971. Fishes of Montana. Montana State University Press,

Bozeman.

3. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

4. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

5. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor. 587

6. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

7. Reisman, H.M., and T.J. Cade. 1967. Physiological and Behavioral Aspects of

Reproduction in the Brook Stickleback, Culaea inconstans. The American

Midland Naturalist, 77(2): 257-295.

588

CHAPTER 20

FAMILY MORONIDAE

Introduction

The Temperate Bass family, Moronidae, stems from the order Perciformes, which is a group of perch-like fishes with over 9,000 species including Walleye, Sander vitreus, in the family Percidae, and Largemouth Bass, Micropterus salmoides, from the family

Centrarchidae. Within the order Perciformes, the Temperate Bass family, Moronidae, is rather small and only contains 6 species, with 4 of the species occurring in North

America, and only two species occurring in and native to the Mississippi River basin: the

White Bass, Morone chrysops, and the Yellow Bass, Morone mississippiensis. The White

Bass is the only species of the family Moronidae to occur in the Dakotas. Temperate Bass are defined by having laterally compressed and rather deep bodies with two dorsal fins.

The anterior dorsal fin is spinous, and the posterior dorsal fin has soft rays. The family also displays a small, flat spine on the posterior edge of each gill cover. Temperate Bass are very similar in appearance to members of the sunfish family Centrarchidae (especially the genus Micropterus), and members of the perch family, Percidae. Moronids differ from Centrarchids by having a defined separation or only slight connection between the two dorsal fins, whereas the two dorsal fins are clearly connected in Centrarchids. Percids have 1-2 spines in the anal fin, whereas Moronids have 3 or more.

Temperate Bass are either entirely freshwater or coastal, anadromous species that occur within North America, Europe, and Africa. The White Bass and Yellow Bass are strictly inland species only inhabiting freshwaters. Although the White Bass is native to 589 the Mississippi River basin in the Dakotas, it is non-native to the Missouri River basin.

Temperate Bass, including the White Bass, are often found schooling in the open waters of lakes, reservoirs, and medium to large sized rivers during the summer and winter months. In the spring and fall months, they are often found closer to shore, likely due to feeding and spawning habits. White Bass are opportunistic sight feeders that forage upon zooplankton, aquatic insects, and other smaller species of fish, especially when they reach adulthood. Moronids have small, but well developed sharp teeth present on both jaws that help them consume their prey. Unlike Largemouth and Smallmouth Bass,

Micropterus dolomieu, from the family Percidae, Moronids do not create nest or provide parental care after spawning.

White Bass are an important species of sportfish in the Dakotas region, and also have been stocked in many other lakes and impoundments across the United States for sport fishing. Striped Bass, Morone saxatilis, does not occur in the Dakotas, however where the species coincides with White Bass, they are known to hybridize. These hybrids are called “wipers”.

590

White Bass, Morone chrysops (Rafinesque, 1820)

Etymology and Synonyms: Morone = definition of origin unknown; chrysops = “golden eye”, in reference to the yellowish tint in the eye.

Description: Body deep, laterally compressed with arched dorsal side. Dorsally dark blue-gray to dark silver; laterally silver to pale green with 6-8 dusky to prominent dark horizontal stripes, unbroken above lateral line; ventrally white; pelvic fin white to light gray, remaining fins dark gray to brown; eye with a yellow-gold tint. Head small, conical.

Snout pointed. Eye moderately large, placed laterally on head. Sharp spine present at rear end of opercle. Mouth terminal and oblique; upper jaw extends below eye; lower jaw slightly extends past upper jaw. Teeth small, sharp; present in patches on both jaws; single patch present on posterior end on tongue. Gill rakers 20-25. Dorsal fin with two separate lobes, close together but not joined; anterior fin with 9 spines; posterior dorsal fin with 1 spine and 12-14 rays. Adipose fin absent. Caudal peduncle short, thick. Caudal fin slightly forked. Anal fin with 3 spines in ascending length, with anterior spine being shortest, and 11-13 rays. Pelvic fins thoracic. Pectoral fins with 15-17 rays. Lateral line complete with 50-60 ctenoid scales in series. Spawning adults may develop a light blue tint on lower jaw, otherwise similar to non-spawning adults. Juveniles less deep bodied.

Similar Species: Although not common in the Dakotas, Striped Bass closely resemble

White Bass. Striped Bass have a less deep and more elongate body, 10-12 rays in the posterior dorsal fin, and 55-63 ctenoid scales in the lateral line.

Distribution and Habitat: Native to the Minnesota and Big Sioux River drainages of eastern South Dakota in the west, east through the Mississippi, Hudson Bay, and St.

Lawrence-Great Lakes basins to New York, and south to Lousiana.12 Introduced across 591 the Dakotas including the Missouri River reservoirs and eastern glacial lakes.1 Inhabits the open waters of lakes, reservoirs, and medium to large sized rivers with clear to turbid water with moderate flow. In eastern South Dakota, White Bass were observed to prefer offshore habitat during summer and winter, and shallow waters near shore in the spring and fall, which is likely related to feeding and spawning behaviors.12

Reproduction: Spawning takes place May to June, or when water temperatures reach roughly 14-22°C (57.2-71.6°F) in tributaries or windswept shorelines of lakes and reservoirs with moderate current, often near the surface over rocky substrates.7,8,10 In

Lewis and Clark Lake, South Dakota, females reach sexual maturity at age-4, or at lengths 250-339 mm (9.84-13.34 in) TL, and males at age-3, and lengths 220-312 mm

(8.66-12.28 in) TL.9 No nest is prepared and no parental care is given. A female 390 mm

(15.35 in) TL may produce 429,000-608,000 eggs.10 Fecundity increases with the length of female.10 Females known to spawn roughly 50% of their eggs during the season, with the smaller eggs being reabsorbed later in the summer.10 Eggs roughly 0.7-1.2 mm (0.03-

0.05 in) in diameter, demersal, and adhesive.3 Hatching takes place in roughly 2 days.7

Spawning season may take place for several weeks.5,8 Successful recruitment in eastern

South Dakota glacial lakes is positively influenced by spring precipitation and air temperature.12 Recruitment highly variable within a region.6,12 High levels of discharge and sudden declines in water temperature may negatively influence spawning activity.8,11

Age and Growth: Larvae roughly 2 mm (0.08 in) TL at hatching.3 Growth of age-0

White Bass in Lewis and Clark Lake, South Dakota is positively related to water temperature and abundance of food.9 Mean lengths-at-age from populations across South

Dakota are reported as: age-1, 145 mm (5.71 in) TL; age-2, 243 mm (9.57 in) TL; age-3, 592

298 mm (11.73 in) TL; age-4, 339 mm (13.35 in) TL; age-5, 358 mm (14.09 in) TL; age-

6, 384 mm (15.12 in) TL; age-7, 394 mm (15.50 in) TL; age-8, 403 mm (15.86 in) TL.13

Females tend to grow faster in length and weight than males.6 Growth rates greatest in eastern South Dakota glacial lakes, intermediate in Missouri River reservoirs, and slowest in western reservoirs.13 Capable of reaching 533.4 mm (21 in) TL. Longevity 14 years.12

Food and Feeding: Opportunistic feeders in response to food availability. Age-0 white bass primarily consume zooplankton, especially Daphnia and calanoid copepods, but may also consume small fish such as Fathead Minnows and Johnny Darters.2,4,12 Fish are typically not an important prey source for White Bass until they reach roughly 40 mm

(1.57 in) TL.8,9 In Lewis and Clark Lake, South Dakota, Emerald Shiners and age-0

Gizzard Shad were common in the diet of White Bass 41-80 mm (1.61-3.15 in) TL, and fishes were the most dominant prey item in individuals >90 mm (3.54 in) TL.9 Adults primarily piscivorous often consuming Gizzard Shad, but also consume aquatic macroinvertebrates such as amphipods, corixids, and dipterans.9,12

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Beck, H.D., A.B. Starostka, and D.W. Willis. 1999. Early life history of white

bass in Lake Poinsett. South Dakota Department of Game, Fish and Parks,

Completion Report 99-14.

3. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

4. Blackwell, B.G., C.A. Soupir, and M.L. Brown. 1999. Seasonal diets of walleye

and diet overlap with other top-level predators in two South Dakota lakes. South 593

Dakota Department of Game, Fish and Parks, Fisheries Division Report 99-23,

Pierre.

5. Guy, C.S., R.D. Schultz, and C.A. Cox. 2002a. Variation in gonad development,

growth, and condition of white bass in Fall River Reservoir, Kansas. Journal of

Fisheries Management 22:643-651.

6. Guy, C.S., R.D. Schultz, and M.A. Colvin. 2002b. Ecology and management of

white bass. North American Journal of Fisheries Management 22:606-608.

7. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

8. Quist, M.C., C.S. Guy, and R.J. Bernot. 2002. Ecology of larval white bass in a

large Kansas reservoir. North American Journal of Fisheries Management 22:637-

642.

9. Ruelle, R. 1971. Factors influencing growth of white bass in Lewis and Clark

Lake. Pages 411-423 in G.E. Hall, editor. Reservoir fisheries and limnology.

American Fisheries Society, Special Publication 8, Bethesda, Maryland.

10. Ruelle, R. 1977. Reproductive cycle and fecundity of white bass in Lewis and

Clark Lake. Transactions of the American Fisheries Society 106:67-76.

11. Starnes, L.B., P.A. Hackney, and T.A. McDonough. 1983. Larval fish transport: a

case study of white bass. Transactions of the American Fisheries Society 112:390-

397.

12. Willis, D.W., C.P. Paukert, and B.G. Blackwell. 2002. Biology of white bass in

eastern South Dakota glacial lakes. North American Journal of Fisheries

Management 22:627-636. 594

13. Willis, D.W., H.D. Beck, C.A. Soupir, B.A. Johnson, G.D. Simpson, and G.A.

Wickstrom. 1997. Growth of white bass in South Dakota waters. South Dakota

Department of Game, Fish and Parks, Completion Report 96-16.

595

CHAPTER 21

FAMILY CENTRARCHIDAE

Introduction

The Sunfish family, Centrarchidae, consists of eight genre and more than 30 species, all of which are native to North America, mainly east of the Continental Divide.

Only one species, the Sacremento Perch, Archoplites interruptus, has its native range west of the continental divide in California. With most species highly valued for sport fishing, many centrarchids have been introduced outside of their native ranges throughout

North and South America, Africa, Asia, and Europe, sometimes leading them to become invasive species. Two of the main genera of Centrarchids in the Dakotas (Lepomis and

Micropterus) are well defined by their simple body morphology. Species in the genera

Lepomis have a deep, more round body shape with smaller mouths, whereas species in the genera Micropterus are more elongate and have larger mouths. Centrarchids are often times confused with the families Percidae (perch) and Moronidae (temperate bass). The most distinguishable feature separating the Centrarchids from these two families is the presence of the fused membrane of the spinous and rayed dorsal fins that are only slightly separated by a shallow notch. Percids and Moronids have barely connected or separate spinous and rays dorsal fins. Centrarchids are also distinguished by having ≥ 3 anal spines, whereas Percids only have 1-2. Spawning male sunfish are also known to develop bright coloration on the cheeks and ventral sides during the springtime.

Centrarchids are considered warmwater adapted species and occur in a variety of lentic and lotic habitats such as swamps, ponds, lakes, reservoirs, and areas with minimal 596 flow in small streams and rivers. Centrarchids can be found at various depths throughout the water column, as the preference often depends on the species. For example, Green

Sunfish, Lepomis cyanellus, frequent the littoral zone, whereas Bluegill, Lepomis macrochirus, are more often found within the limnetic zone. They are most often found as individuals or small groups rather than large schools among varieties of partially or fully submerged aquatic and overhanging vegetation which serve as cover from predators or as a place to hide from potential prey items. Centrarchids primarily feed by sight during the day, and most food is consumed by active foraging or ambush behavior.

Where feeding occurs is species specific, but Centrarchids are known to consume their prey items from the surface, within the water column, along vegetation and submerged structures, as well as along the bottom substrate. Aquatic and terrestrial invertebrates as well as small fish are common components of Centrarchid diets, however foraging behavior is known to be heavily influenced by other species interactions within the same habitat.

Spawning behavior of Centrarchids is rather intricate, as most members within the family exhibit courtship, nest construction, and parental care. Males of certain species will excavate a saucer shaped nest using their fins within the substrate. The nests may be constructed within large colonies, or be more dispersed within a spawning site. After the eggs are deposited in the nest by the female, most Centrarchid males will provide parental care by fanning the eggs, and guarding the eggs and fry pre- and post- hatching.

Because many species of Centrarchidae exhibit very similar spawning behavior, especially in Lepomis species, hybridization often occurs. 597

In addition to their current occurance throughout the United States, crappies,

Pomoxis sp., larger Lepomis sp., basses, Micropterus sp., and the Rock Bass, Ambloplites rupestris, are all important game fishes of the Dakotas. In many lakes across the states, populations are density dependent where overpopulation may cause stunting in growth.

598

Rock Bass, Ambloplites rupestris (Rafinesque, 1817)

Etymology and Synonyms: Ambloplites = Greek, amblo-, meaning “blunt”, and –plites, meaning “armature”; rupestris = “living among rocks”, referring to the preferred habitat of the species.

Description: Body deep, robust, laterally compressed. Dorsally dark brown to olive; laterally solid olive to bronze, but may display dark mottling and dark spots on scales below lateral line forming 8-10 horizontal rows; ventrally tan to creamy white; dorsal, caudal, and anal fins with dark and light mottling; pelvic and pectoral fins dusky. Capable of changing coloration to camouflage with habitat. Head large, robust. Snout conical. Eye large, partially dark amber-brown in color, placed laterally on head. Mouth large, terminal, and oblique; upper jaw extends to or beyond middle of pupil; lower jaw protrudes past upper jaw. Teeth small, blunt, present in pads on upper and lower jaws; tooth patch present on tongue. Gill rakers long, thin, 7-10. Dorsal fin with two lobes broadly joined appearing as one; 10-12 spines followed by 10-12 rays. Adipose fin absent. Caudal peduncle thick, slightly elongate. Caudal fin slightly forked. Anal fin with

5-7 spines and 9-11 rays. Pelvic fin with 1 spine and 5 rays. Pectoral fin short, rounded; does not extend to eye when folded forward. Lateral line complete, arched upward with

37-46 ctenoid scales in series. Spawning males with darker coloring on body, and edges of anal and pelvic fins; iris becomes red. Juveniles more likely to be mottled than adults.

Similar Species: Does not closely resemble any other species within the Dakotas. Differs from other sunfish species within the Dakotas by having a greater number of spines in the dorsal and anal fins. Largemouth and Smallmouth Bass with 9-10 spines in dorsal fin. 599

Distribution and Habitat: Native to North America from Saskatchewan and eastern

North and South Dakota in the west, east throughout the Upper Mississippi, Great Lakes, and Hudson Bay drainages to Quebec and Vermont, and south to the Savannah River drainage and northern Alabama. Occurs throughout the Missouri and Red River drainages in the Dakotas, with distant populations present in Sheridan Reservoir and small lakes in the Black Hills of South Dakota. Most frequent in cool to warm waters of rivers, streams, lakes, and reservoirs in clear to slightly turbid water with slow current, dense aquatic vegetation, submerged structure, and rocky substrate. In lotic habitats, areas with slow current such as pools and backwaters are preferred. Smaller individuals utilize shallow and quiet areas more frequently than large individuals.4 Overwinter in deeper waters away from shorelines.1

Reproduction: Spawning takes place May to July. Females have shown peak spawning behavior at 21-23°C (69.8-73.4°F).6 Sexual maturity reached at age 3-4 in stream populations and age 5-9 in lakes.9 Males become territorial while preparing saucer- shaped nests roughly 20-25 cm (8-10 in) that are excavated in coarse substrate using the anal and pectoral fins in water 0.5-1.5 m (1.64-4.92 ft) deep in close proximity to cover.6,8,9 Nest construction begins when water temperature reaches approximately 20°C

(68°F).1,11 Spawning ritual typically takes place 2 days after nest construction.6 Females enter nests without being courted by males.6 Spawning occurs in shallow waters over gravel substrate and is most frequent in the early mornings or late evenings.6 Males become darker in color during the spawning act.6 Fractional spawners. Fecundity ranges

3,000-11,000 eggs per female. Mature eggs orange in color, roughly 1.7-1.9 mm (0.06-

0.07 in) in diameter and adhesive.1 Males provide parental care by fanning the eggs and 600 defending them from predators until larvae leave the nest in 9-10 days post hatching.6

Hatching occurs within 5 days in water 16-22°C (60.8-71.6°F).6 Spawning known to cease when water temperatures reach 26°C (78.8°F).1,11

Age and Growth: Growth variable among location and populations. Larvae average 5.6 mm (0.22 in) TL at time of hatching.2 Mean lengths at age from Missouri were reported as: age-1, 41 mm (1.61 in) TL; age-2, 86 mm (3.39 in) TL; age-3, 140 mm (5.51 in) TL; age-4, 178 mm (7.00 in) TL; age-5, 203 mm (7.99 in) TL; age-6, 216 mm (8.50 in) TL.10

Capable of reaching 431.8 mm (17 in) TL.10 Average lifespan 6-8 years; longevity 10 years.1,10

Food and Feeding: Generalist predator. Young-of-year feed on ephemeroptera, cladocerans, copepods, chironomid larvae, amphipods, and isopods.5,7 Individuals 150-

200 mm (5.91-7.87 in) TL mainly consume snails and chironomids.3 Adults mainly consume amphipods, trichopterans, gastropods, decapods, chironomids, and minnows.3,4

As mean length increases, decapods and fish increase in the diet and amphipods and trichopterans occur less frequently.3

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Buynak, G.L., and H.W. Mohr Jr. 1979. Larval development of rock bass from

the Susquehanna River. The Progressive Fish Culturist 41:39-42.

3. Elrod, J.H., W.D.N. Busch, B.L. Griswold, C.P. Schneider, and D.R. Wolfert.

1981. Food of white perch, rock bass and yellow perch in eastern Lake Ontario.

New York Fish and Game Journal 28:191-201. 601

4. George, E.L., and W.F. Hadley. 1979. Food and habitat partitioning between rock

bass (Ambloplites rupestris) and Smallmouth Bass (Micropterus dolomieui)

young of the year. Transactions of the American Fisheries Society 108:253-261.

5. Glessner, G.L. 1977. Food habits and growth of the rock bass, Ambloplites

rupestris (Rafinesque), in Stone Valley Lake, Huntingdon County, Pennsylvania.

M.S. thesis. Pennsylvania State University, University Park, Pennsylvania.

6. Gross, M.R., and W.A. Nowell. 1980. The reproductive biology of rock bass,

Ambloplites rupestris (Centrarchidae), in Lake Opinicon, Ontario. Copeia 3:482-

494.

7. Keast, A. 1977a. Mechanisms minimizing intraspecific competition in vertebrates,

with a quantitative study of the contrasting strategies of two centrarchid fishes,

Ambloplites rupestris and Lepomis macrochirus. Evolutionary Biology 10:333-

395.

8. Musch, A.E. 2007. Spawning habitat selection of sympatric smallmouth bass

(Micropterus dolomieu) and Rock Bass (Ambloplites rupestris) in north temperate

lakes: habitat separation in space and time. M.S. thesis, University of Wisconsin-

Stevens Point.

9. Noltie, D.B., and M.H.A. Keenleyside. 1986. Breeding ecology, nest

characteristics, and nest-site selection of stream- and lake-dwelling rock bass,

Ambloplites rupestris (Rafinesque). Canadian Journal of Zoology 65:379-390.

10. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 602

11. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

603

Green Sunfish, Lepomis cyanellus (Rafinesque, 1819)

Etymology and Synonyms: Lepomis = Greek, for “scaled lid”, referring to the scaled operculum; cyanellus = “blue”.

Description: Body deep, robust, slightly elongate, laterally compressed, with arched dorsal profile. Dorsally dark olive to emerald green with emerald reflections; laterally green to light green with blue-green reflections and 7-12 extremely faint, dark vertical bars; ventrally yellow to cream; fins uniformly pigmented with dark blotch present on posterior base of dorsal fin; head with backward extending wavy light blue-green lines or spots; opercle with a dark center and light border. Head and snout large, conical. Eye moderately large, placed laterally on head. Mouth terminal, slightly oblique, large; upper jaw extends to middle of eye. Lips fleshy, smooth. Teeth small, present in pads on both jaws; absent or little on tongue. Gill rakers long, slender, 11-14 on first arch. Opercular flap elongate, inflexible. Dorsal fin elongate with 9-11 spines followed by 10-11 rays.

Adipose fin absent. Caudal peduncle short, thick. Caudal fin slightly forked with rounded lobes. Anal fin with 3 spines and 8-10 rays. Pelvic fin with 1 spine and 5 rays. Pectoral fin rounded on distal end with 13-15 rays; barely reaches eye when folded forward.

Lateral line complete, arched upward with 44-53 ctenoid scales in series. Juveniles similar to adults. Spawning males with more intense coloration with distinct yellow-white margin on distal end of dorsal, caudal, and anal fins.

Similar Species: Closely resembles other Lepomis species within the Dakotas. Bluegill have a deeper, less elongate body shape, a short, flexible solid blue opercle, and have an anal fin with 10-12 rays. Pumpkinseed are easily distinguished by the single red spot on the posterior edge of the opercle. Orangespotted Sunfish display small, irregular orange 604 spots along the lateral sides, as well as a single pair of large sensory pores between the eyes.

Distribution and Habitat: Native to central North America from southwest South

Dakota, east throughout the Missouri, Mississippi, and Great Lakes drainages to eastern

Pennsylvania, and south to Alabama and eastern New Mexico. Widely introduced and dispersed outside its native range due to its ability to colonize, withstand, and exploit new habitats after flooding events. Occurs throughout the major tributaries of the Missouri

River in the Dakotas. Most abundant in low gradient, lacustrine environments including ponds, lakes, and reservoirs with large littoral areas, abundant submerged vegetation, sand, gravel or silt substrates, and moderate turbidity.14 Also inhabits low flow areas such as pools in small streams and rivers. Optimal water temperature 28.2°C (82.8°F) and avoids temperature ˃31°C (87.8°F), and ˂26°C (78.8°F).2,16 Mean critical thermal maximum 36.5-37.9°C (98-100°F).15 Tolerant of high turbidity, high alkalinity, low dissolved oxygen levels, and rapid changes in pH. Highly turbid water known to decrease activity levels.7

Reproduction: Spawning takes place May to early July, or when water temperatures reach 19-28°C (66.2-82.4°F). Sexual maturity reached at age 1-3, depending on geographic range. Males prepare and defend nests roughly 30 cm (11.8 in) in diameter in colonies or singularly near vegetative cover or large rocks in depths of 4-35 cm (0.13-1.4 ft) in sand or gravel substrate.4,8 Spawning begins 1-2 days following nest construction.8

Males court females with a series of grunting sounds or visual cues.5 Males provide parental care for roughly a week while eggs are developing and fry flee the nest.8 Eggs 605 roughly 1.0 mm (0.04 in) in diameter, demersal, and adhesive. Hatching occurs within

35-55 hours in water temperatures of 24-27°C (75.2-80.6°F).17

Age and Growth: Newly hatched larvae 3.5-3.7 mm (0.13-0.14 in) TL.17 Growth rates highly variable between populations, but known to increase with density of woody debris.12 Stunting may occur in overpopulated, small closed systems. Capable of reaching

304.8 mm (12 in) TL. Average life span 5-6 years; longevity 10 years.4

Food and Feeding: Voracious predator. Feeds by sight, and uses lateral line system to detect prey in turbid waters.9 Feeding takes place day and night for fry and young ≤9.5 mm (0.37 in) TL.1 As they grow, foraging is more intense during the day.1 Fry initially consume zooplankton, and add immature aquatic insects and egg cases as they grow.13 In

South Dakota, fry primarily selected for the small crustaceans Cyclops vernalis and

Moina brachiata.1 Mean length of prey increases with the mean length of Green Sunfish fry.1 Adults primarily feed on insects, crayfish, and small fish.11 Reduced competition increases average prey size and growth.18 Juvenile diet similar to adults. Juveniles possess the ability to detect chemical alarm signals from other Green Sunfish when their skin cells become damaged from predators.3,6 When exposed to the chemical alarm signals, small individuals exhibit fin-erect posture, while larger individuals may continue to move as if it were foraging.3,6,10

Literature Cited:

1. Barkoh, A. 1984. Food selectivity of bluegill and green sunfish fry. M.S. Thesis,

South Dakota State University, Brookings. 606

2. Beitinger, T.L., J.J. Magnuson, W.H. Neill, and W.R. Shaffer. 1975. Behavioral

thermoregulation and activity patterns in the green sunfish, Lepomis cyanellus.

Animal Behavior 23;222-229.

3. Brown, G.E., and S. Brennan. 2000. Chemical alarm signals in juvenile green

sunfish (Lepomis cyanellus, Centrarchidae). Copeia 4:1079-1082.

4. Carlander, K.D. 1977. Handbook of freshwater fishery biology, Vol. 2. Iowa State

University Press, Ames. 431pp.

5. Gerald, J.W. 1971. Sound production during courtship in six species of sunfish

(Centrarchidae). Evolution 25:75-87.

6. Golub, J.L., and G.E. Brown. 2003. Are all signals the same? Ontogenetic change

in the response to conspecific and heterospecific chemical alarm signals by

juvenile green sunfish (Lepomis cyanellus). Behavioral Ecology and

Sociobiology 54:113-118.

7. Horkel, J.D., and W.D. Pearson. 1976. Effects of turbidity on ventilation rates and

oxygen consumption of Green Sunfish, Lepomis cyanellus. Transactions of the

American Fisheries Society 105:107-113.

8. Hunter, J.R. 1963. The reproductive behavior of the green sunfish, Lepomis

cyanellus. Zoologica 48:13-24.

9. Janssen, J., and J. Corcoran. 1993. Lateral line stimuli can override vision to

determine sunfish strike trajectory. Journal of Experimental Biology 176:299-305.

10. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence. 607

11. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

12. Quist, M.C., and C.S. Guy. Growth and mortality of prairie stream fishes:

relations with fish community and instream habitat characteristics. Ecology of

Freshwater Fish 10:88-96.

13. Sadzikowski, M.R., and D.C. Wallace. 1976. A comparison of the food habits of

size classes of three sunfishes (Lepomis macrochirus Rafinesque, L. gibbosus

(Linnaeus) and L. cyanellus Rafinesque). American Midland Naturalist 95:220-

225.

14. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

15. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

16. Stuber, R.J., G. Gebhart, and O.E. Maughan. 1982. Habitat suitability index

models: Green sunfish. U.S. Department of Interior Fish and Wildlife Service.

FWS/OBS-82/10.15. 28pp.

17. Taubert, B.D. 1977. Early morphological development of the green sunfish,

Lepomis cyanellus, and its separation from other larval Lepomis species.

Transactions of the American Fisheries Society 106:445-448.

18. Werner, C.A., and D.J. Hall. 1976. Niche shifts in sunfishes: experimental

evidence and significance. Science 191:404-406.

608

Pumpkinseed, Lepomis gibbosus (Linnaeus, 1758)

Etymology and Synonyms: Lepomis = Greek, lepis- meaning “scaled”, and –poma meaning “lid”, meaning scaled operculum; gibbosus = “formed like the full moon”, likely referring to the body shape.

Description: Body deep, laterally compressed with arched dorsal profile. Dorsally golden-brown to olive; laterally tan with 7-10 faint, dark, diffuse vertical bars (most prominent in females) with orange, emerald, or yellow reflections; ventrally yellow- orange anteriorly fading to light yellow-cream posteriorly; dorsal, caudal, and anal fins dark with faint to prominent small, brown spots or blotches; head with 3-5 wavy blue- green lines extending backward to gill flap; opercle with a dark black center and white cream margin separated by single red spot on posterior edge. Head and snout large, conical. Eye moderately large, placed laterally on head. Mouth terminal, slightly oblique, small; upper jaw barely extends to anterior end of eye; lower jaw slightly protrudes past upper jaw. Lips fleshy. Teeth small, present in pads on both jaws; absent on tongue. Gill rakers short, thick, 9-12. Opercular flap short, flexible. Dorsal fin elongate with 10-11 spines followed by 10-12 rays. Adipose fin absent. Caudal peduncle slightly elongate, thick. Caudal fin slightly forked with rounded lobes. Anal fin with 3 spines and 9-10 rays.

Pelvic fin with 1 spine and 5 rays. Pectoral fin with pointed distal end and 12-14 rays; extends to anterior end of eye when folded forward. Lateral line complete, arched upward with 36-43 ctenoid scales in series. Juveniles similar to adults with less intense and dull coloration. Spawning males with intense bright coloration, especially on throat region and red spot on posterior edge of opercle. 609

Similar Species: Closely resembles other Lepomis species within the Dakotas. Easily distinguished by the single red spot on posterior edge of opercle. Redear Sunfish lack any blue-green way lines on head, and have a short and stiff opercle flap with a black center, and a red to burnt-orange, crescent-shaped margin on the posterior edge. Orangespotted

Sunfish display orange spots rather than reflections on the lateral sides, have large sensory pores present between the eyes, and are typically smaller in size. Young-of-year

Pumpkinseed may resemble the Bluegill; however Bluegill display a solid dark blue opercle. Green Sunfish have a more elongate body and are often dark in color with 44-53 scales in lateral line series.

Distribution and Habitat: Native to the northeastern United States from eastern North and South Dakota and southeastern Manitoba in the west, east into the upper Mississippi

River basin, Great Lakes, and Hudson Bay drainages to New England territory, and south through the Atlantic Slope drainages to South Carolina. Widely introduced and distributed. Stocked variously and occurs throughout the main tributaries of the Missouri

River in the Dakotas. Inhabits clear, quiet waters of ponds, lakes, and reservoirs, but can also occur in low flow areas of small streams and rivers. Often associated with abundant aquatic vegetation. Actively swims in the midwater.8 Tolerant of a wide variety of environmental conditions including low dissolved oxygen and moderate turbidity. Less tolerant of warmer water temperature than Bluegill.10 Preferred water temperature range

24-32°C (75.2-89.6°F).5 Home ranges vary from 0.23-1.12 ha (0.57-2.77 ac).6

Reproduction: Spawning takes place May to August, or when water temperatures near

20°C (68°F).1,8,12 Mean age of sexual maturity in males is 3.5 years, or 104 mm (4.09 in)

TL; females 3.4 years and 100 mm (3.94 in) TL.7 Larger individuals mature earlier than 610 smaller individuals in the same population of the same age.7 Males excavate and defend saucer-shaped nests, roughly twice the length of the male in diameter.8 Nests often constructed singularly or in colonies near submerged structure in quiet shallow areas ˂1 m (3.3 ft) in depth with partial sunlight, little to no aquatic vegetation, and sand, gravel, or mud substrate.8 Females remain in deeper water and wait to move into the nesting site until nest construction is completed.8 Fractional spawners; females spawn in more than one nest throughout the season. Males provide parental care by vigorously defending and fanning the eggs until hatching occurs.8 Fecundity dependent on size and age of female, but may reach 7,000 eggs per female.1 Eggs creamy amber-white in color, roughly 1.0 mm (0.04 in) in diameter, demersal and adhesive.1,8 Hatching occurs within 2-3 days at a water temperature of 27.8°C (82°F), and may be longer depending on temperature.2

Age and Growth: Growth highly variable among populations depending on density, temperature, and water quality. Stunting occurs frequently in populations of high density.

Average length 127-203 mm (5-8 in) TL but capable of reaching 254 mm (10 in) TL.4

Longevity 8-10 years.

Food and Feeding: Primarily feeds on benthic invertebrates such as aquatic insects and larvae, snails, small crustaceans, and small fish.8,9 Ontogenetic diet shift from littoral prey to open water prey occurs between 45-70 mm (1.77-2.76 in) SL.9 Pharyngeal jaws assist in crushing mollusks, and have been shown to be morphologically different in lakes with and without an abundance of snails.13 Also known to forage on large proportions of invasive zebra mussels during spring and summer.3 Juveniles are the most food limited in lakes with Blugill.11 Adults are most food limited in lakes without Bluegill.11

Literature Cited: 611

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Breder, C.M. 1936. The reproductive habits of the North Amercian sunfishes

(family Centrarchidae). Zoologica 21:1-48.

3. Colborne, S.F., A.D.M. Clapp, F.J. Longstaffe, and B.D. Neff. 2015. Foraging

ecology of native pumpkinseed (Lepomis gibbosus) following the invasion of

zebra mussels (Dreissena polymorpha). Canadian Journal of Fisheries and

Aquatic Sciences 72:983-990.

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

5. Evans, D.O. 1977. Seasonal changes in standard metabolism, upper and lower

thermal tolerance and thermoregulatory behavior of the pumpkinseed, Lepomis

gibbosus, Linnaeus. Doctoral dissertation. University of Toronto, Toronto,

Canada.

6. Fish, P.A., and J. Savitz. 1983. Variations in home ranges of largemouth bass,

yellow perch, bluegills, and pumpkinseeds in an Illinois lake. Transactions of the

American Fisheries Society 112:147-153.

7. Fox, M.G. 1994. Growth, density, and interspecific influences on pumpkinseed

sunfish life histories. Ecology 75:1157-1171.

8. Miller, H.C. 1963. The behavior of the pumpkinseed sunfish, Lepomis gibbosus

(Linneaus), with notes on the behavior of other species of Lepomis and the pigmy

sunfish, Elassoma evergladei. Behavior 22:88-151.

9. Mittelbach, G.G. 1984. Predation and resource partitioning in two sunfishes

(Centrarchidae). Ecology 65:499-513. 612

10. O’Hara, J.J. 1968. Influence of weight and temperature on metabolic rate of

sunfish. Ecology 49:159-161.

11. Osenberg, C.W., G.G. Mittelbach, and P.C. Wainwright. 1992. Two-stage life

histories in fish: the interaction between juvenile competition and adult

performance. Ecology 73:255-267.

12. Reed, R.J. 1971. Underwater observations of the population density and behavior

of pumpkinseed, Lepomis gibbosus (Linnaeus) in Cranberry Pond, Massachusetts.

Transactions of the American Fisheries Society 100:350-353.

13. Wainwright, P.C., C.W. Osenberg, and G.G. Mittelbach. 1991. Trophic

polymorphism in the pumpkinseed sunfish (Lepomis gibbosus Linnaeus): effects

of environment on ontogeny. Functional Ecology 5:40-55.

613

Orangespotted Sunfish, Lepomis humilis (Girard, 1858)

Etymology and Synonyms: Lepomis = Greek, lepis- meaning “scaled”, and –poma meaning “lid”, meaning scaled operculum; humilis = “humble” or “insignificant”, possibly referring to the species small size.

Description: Body moderately deep, laterally compressed, with arched dorsal profile.

Dorsally olive to blue-brown; laterally bluish gray to blue-green with small, irregular orange spots and 5-7 extremely faint, dark vertical bands; ventrally white, yellow, or light orange; dorsal, caudal, and anal fins dark with small, faint to dark blotches or spots present, often near base; lower jaw with occasional backward extending wavy blue lines under eye; opercle with a dark center and light border. Head and snout large, conical. Eye moderately large, placed laterally on head. Single pair of large sensory pores present between eyes above upper lip. Mouth terminal, oblique, moderately large; upper jaw extends to or slightly past middle of eye; lower jaw slightly protrudes past upper jaw.

Lips fleshy. Teeth small, present in pads on both jaws; absent on tongue. Gill rakers long, thin, 10-15. Opercular flap short, flexible. Dorsal fin elongate with 10-11 spines followed by 9-10 rays. Adipose fin absent. Caudal peduncle slightly elongate, thick. Caudal fin slightly forked with rounded lobes. Anal fin with 3 spines and 8-9 rays. Pelvic fin with 1 spine and 5 rays. Pectoral fin slightly pointed with 14-15 rays; extends to or barely reaches anterior end of eye when folded forward. Lateral line complete, arched upward with 32-41 ctenoid scales in series. Juveniles similar to adults with less intense coloration and spots, and more prominent, dusky vertical bars on lateral sides. Spawning males with more intense coloration, especially on fins, lateral, and ventral sides; pelvic and anal fins develop a black border on distal end; eye becomes red-orange in color. 614

Similar Species: Closely resembles other Lepomis species within the Dakotas. Easily distinguished by the large sensory pores between the eyes. Green Sunfish lack orange spots on lateral sides, are much darker in color, more elongate, and have 44-53 scales in lateral line series. Pumpkinseed display a single red spot on posterior edge of opercle.

Bluegill have a solid dark blue opercle without a light border.

Distribution and Habitat: Native to the central United States from the Missouri River basin in eastern North and South Dakota in the west, east throughout the Mississippi

River basin and Great Lakes drainages to Ohio, and south to the Gulf of Mexico from

Alabama to Texas. Most abundant east of the Missouri River in the Dakotas. Most frequent in low flow areas such as pools and backwaters in small to large streams hovering above mud, gravel, or sand substrate. Less abundant in ponds and lakes than other species of sunfish. Tolerant of siltation, high water temperatures and low dissolved oxygen levels. Mean critical thermal maximum reported as 37.9°C (100.22°F).7 Mean critical dissolved oxygen minimum reported as 0.63 mg/L.7

Reproduction: Spawning activity initiated when water temperature reaches 18°C

(64.4°F).1,5 Spawning takes place May to July, or when water temperatures reach 24-

32°C (75-89°F).2 Sexual maturity reached at age 2-3. Males excavate and defend saucer- shaped nests, roughly 15-18 cm (5.91-7.09 in) in diameter and 3-4 cm (1.18-1.57 in) deep, much smaller than other species of sunfish.1,5 Nests often constructed in colonies in shallow, quiet areas with silt, sand, or gravel substrate. Males court females with a series of species-specific grunting sounds.3 Males provide parental care by defending and fanning the eggs free of silt until hatching occurs.1 Darters and minnows known to prey upon eggs while males defend off predators.1,2 Fecundity increases with size of female; a 615

105 mm (4.13 in) TL female produced 4,700 eggs.1 Eggs amber in color, transparent, slightly adhesive, and roughly 0.5-1.0 mm (0.02-0.04 in) in diameter.1 Hatching occurs within 5 days at water temperatures of 18-21°C (64.4-69.8°F).1

Age and Growth: Individuals from the Big Sioux River basin in eastern South Dakota had a mean length of 54 mm (2.13 in) TL with individuals 50-60 mm (1.97-2.36 in) TL dominating the population.6 Individuals from the Cheyenne River basin in western South

Dakota had a mean length of 58 mm (2.28 in) TL with individuals 60-70 mm (2.36-2.76 in) TL dominating the population.6 Mean lengths at age are similar between the Big Sioux and Cheyenne River basins, however all age groups in the Big Sioux are consistently greater.6 Mean lengths-at-age from the Big Sioux River basin are reported as: age-1, 49 mm (1.93 in) TL; age-2, 54 mm (2.13 in) TL; age-3, 61 mm (2.40 in) TL; age-4, 73 mm

(2.87 in) TL.6 Capable of reaching lengths ˃120 mm (4.72 in) TL. Average lifespan 3-4 years; longevity 7 years.1

Food and Feeding: Generalist predator. Consume prey within a wide variety of sizes due to the species small body paired with a large mouth.4 Adults manly consume aquatic invertebrates including midge larvae and small crustaceans. Diet and size of prey known to influence morphological plasticity.4 Age-0 individuals foraging on tiny, planktonic prey developed a long, fusiform body shape and a sharply angled anout.4 Individuals foraging on larger prey developed deeper bodies and a blunt snout.4

Literature Cited:

1. Barney, R.L. and B.J. Anson. 1923. Life history and ecology of the orangespotted

sunfish (Lepomis humilis). Appendix XV, Report of the U.S. Commissioner of

Fisheries for 1922:1-16. 616

2. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

3. Gerald, J.W. 1971. Sound production during courtship in six species of sunfish

(Centrarchidae). Evolution 25:75-87.

4. Hegrenes, S. 2001. Diet-induced phenotypic plasticity of feeding morphology in

the orangespotted sunfish, Lepomis humilis. Ecology of Freshwater Fish 10:35-42.

5. Miller, H.C. 1963. The behavior of the pumpkinseed sunfish, Lepomis gibbosus

(Linneaus), with notes on the behavior of other species of Lepomis and the pigmy

sunfish, Elassoma evergladei. Behavior 22:88-151.

6. Rasmus, R.A., Q.E. Phelps, J.P. Duehr, and C.R. Berry Jr. 2008. Population

characteristics of lotic orangespotted sunfish. Journal of Freshwater Ecology

23:459-461.

7. Smale, M.A., and C.F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of

headwater stream fishes. Transactions of the American Fisheries Society 124:698-

710.

617

Bluegill, Lepomis macrochirus (Rafinesque, 1819)

Etymology and Synonyms: Lepomis = Greek, for “scaled lid”, referring to the scaled operculum; macrochirus = macro- meaning “long”, and –cheir meaning “hand”, in reference to the size of the pectoral fin.

Description: Body deep, robust, strongly laterally compressed, with arched dorsal profile. Dorsally dark olive brown; laterally olive-brown to dark blue-brown sometimes with small blue-purple reflections, and 7-10 faint, dark vertical bars; ventral and throat region white, yellow, or light orange-red; fins uniformly pigmented with dark blotch present on posterior base of dorsal fin and occasionally anal fin; lower jaw with backward extending wavy pale blue lines; opercle solid dark blue. Coloration varies with habitat.

Head and snout large, conical. Eye moderately large, placed laterally on head. Mouth strongly oblique, small; upper jaw does not extend to anterior end of eye. Lips fleshy, smooth. Teeth small, present in pads on both jaws; typically absent on tongue. Gill rakers long, slender, 13-16. Opercular flap short, flexible. Dorsal fin elongate with 9-11 spines and 10-12 rays. Adipose fin absent. Caudal peduncle slightly elongate, thick. Caudal fin slightly forked with rounded lobes. Anal fin with 3 spines and 10-12 rays. Pelvic fin with

1 spine and 5 rays. Pectoral fin long, pointed with 13-14 rays; extends to or just past eye when folded forward. Lateral line complete, arched upward with 39-44 ctenoid scales in series. Juveniles similar to adults with less intense coloration and vertical bars on lateral sides absent. Spawning males with intense bright orange coloration, especially on lateral and ventral sides.

Similar Species: Resembles other Lepomis species in the Dakotas. Green Sunfish have a more elongate body, display an inflexible opercle with a dark center surrounded by a 618 light border, and an anal fin with 8-10 rays. Pumpkinseed and Redear Sunfish display a single red spot on posterior edge of opercle. Orangespotted Sunfish have small, irregular orange spots along the lateral sides, and large sensory pores between the eyes.

Distribution and Habitat: Native to the eastern and central United States, from eastern

South Dakota in the west, east throughout the Great Lakes drainages to New York, and south to the Gulf of Mexico. Widely introduced and established across the United States due to popularity as a sportfish, as well as a prey fish for other sportfish such as

Largemouth Bass.6 Habitat use, home ranges, and movement likely dependent on prey availability and size, vegetation density, avoidance of predators, and interspecific competition.9,10,14 Often associated with complex structural habitats such as submerged woody debris and dense aquatic vegetation across all seasons, with exception to spawning habitat.14,18 Habitat use similar between diel periods.12 Adults known to show no preference between vegetated and open-water habitat, likely due to the fact that they are less susceptible to predation and are able to utilize more habitat types.12

Reproduction: Spawning takes place in water temperature ≥20°C (68°F). Fractional spawner. Sexually mature at age 2-3. Males excavate and defend nests roughly 30 cm

(11.81 in) in diameter in colonies up to 50 nests, within an average depth of 1.0 m (3.28 ft) along shorelines with gravel or sand substrate, low densities of submerged vegetation, and moderate levels of dissolved oxygen.4,18 Males court females with calls, or grunts.3

Smaller, faster maturing males called “sneakers” may sneak into a larger, older males nest to fertilize with the female.5 Males provide parental care by fanning and guarding the eggs. Eggs small, demersal, and adhesive. After spawning, adults likely move to depths

≥5 m (16.40 ft) to feed on large, open-water zooplankton.9,10,18 Hatching occurs within 2 619 days. Following yolk absorption, larvae migrate to the limnetic zone and then retreat to the littoral zone once they have reached a larger body size.1,19

Age and Growth: Growth affected by water quality, lake morphometry, and temperature.16,15,17 Many populations become stunted and have poor population size structure due to high recruitment rates and lack of predators. Growth rates faster in lakes and impoundments than streams. Deep, large lakes are less optimal habitat for producing fast-growing or strong year classes.16 Predators such as Yellow Perch, Walleye, and

Black or Brown Bullhead often decrease Bluegill densities, consequently increasing

Bluegill growth.17 A high proportion of Bluegills 150 mm (5.90 in) TL and longer can be created in small northern impoundments from a high-density population of Largemouth

Bass shorter than 300 mm (11.81 in) TL, similar to small southern impoundments.6

Growth highly variable between populations. Statewide mean lengths-at-age from South

Dakota were reported as: age-1, 55 mm (2.17 in) TL; age-2, 103 mm (4.06 in) TL; age-3,

141 mm (5.55 in) TL; age-4, 166 mm (6.54 in) TL; age-5, 180 mm (7.09 in) TL.20

Capable of reaching 330 mm (13 in) TL. Longevity 10 years.

Food and Feeding: Generalist, sight feeder. Feeds diurnally, but also observed foraging at night.13 Optimum feeding temperature 27-31°C (80.6-87.8°F).8 Young feed on zooplankton. Adults mainly consume zooplankton such as Daphnia, macroinvertebrates including chironomids, amphipods, trichopterans, and gastropods, and small fish.2,7,9,11

Lakes with high vegetation abundance may increase macroinvertebrate abundance, allowing Bluegill to rely more on macroinvertebrates.11 Zooplankton more likely to be consumed in open water habitat when foraging efficiency is greater and predation risk is limited.9 620

Literature Cited:

1. Dimond, W.F., T.W. Storck, and K.C. Kruse. 1985. A turbidity-related delay in

bluegill (Lepomis macrochirus) reproduction and some size dependent differences

in the spatial distribution of bluegill fry. Transactions of the Illinois State

Academy of Science 78:49-56.

2. Engel, S. 1988. The role and interaction of submersed macrophytes in a shallow

Wisconsin lake. Journal of Freshwater Ecology 4:329-341.

3. Gerald, J.W. 1971. Sound production during courtship in six species of sunfish

(Centrarchidae). Evolution 25:75-87.

4. Gosch, N.J.C., Q.E. Phelps, and D.W. Willis. 2006. Habitat characteristics at

bluegill spawning colonies in a South Dakota glacial lake. Ecology of Freshwater

Fish 15:464-469.

5. Gross, M.R., and W.A. Charnov. 1980. Alternative male life history in bluegill

sunfish. Proceedings of the National Academy of Science USA 77:6937-6940.

6. Guy, C.S., and D.W. Willis. 1990. Structural relationships of largemouth bass and

bluegill populations in South Dakota ponds. North American Journal of Fisheries

Management 10:338-343.

7. Harris, N.J., G.F. Galinat, and D.W. Willis. 1999. Seasonal food habits of

bluegills in Richmond Lake, South Dakota. Proceedings of the South Dakota

Academy of Science 78:79-85.

8. Kitchell, J.F., J.F. Koonce, R.V. O’Neill, H.H. Jr. Shugart, J.J. Magnuson, and

R.S. Booth. 1974. Model of fish biomass dynamics. Transactions of the American

Fisheries Society 103:786-798. 621

9. Mittelbach, G.G. 1981. Foraging efficiency and body size: a study of optimal diet

and habitat use by bluegills. Ecology 62:1370-1386.

10. Mittelbach, G.G. 1984. Predation and resource partitioning in two sunfishes

(Centrarchidae). Ecology 65:499-513.

11. Olson, N.W., C.P. Paukert, and D.W. Willis. 2003. Prey selection and diets of

bluegill Lepomis macrochirus with differing population characteristics in two

Nebraska natural lakes. Fisheries Management and Ecology 10:31-40.

12. Paukert, C.P., and D.W. Willis. 2002. Seasonal and diel habitat selection by

bluegills in a shallow natural lake. Transactions of the American Fisheries Society

131:1131-1139.

13. Sarker, A.L. 1977. Feeding ecology of the bluegill, Lepomis macrochirus, in two

heated reservoirs of Texas III. Time of day and patterns of feeding. Transactions

of the American Fisheries Society 106:596-601.

14. Savino, J.F., and R.A. Stein. 1982. Predator-prey interaction between largemouth

bass and bluegills as influenced by simulated submersed vegetation. Transactions

of the American Fisheries Society 111:255-266.

15. Tomcko, C.M., and R.B. Pierce. 1997. Bluegill growth rates in Minnesota.

Minnesota Department of Natural Resources, Section of Fisheries, Investigational

Report 458, St. Paul.

16. Tomcko, C.M., and R.B. Pierce. 2001. The relationship of bluegill growth, lake

morphometry, and water quality in Minnesota. Transactions of the American

Fisheries Society 130:317-321. 622

17. Tomcko, C.M., and R.B. Pierce. 2005. Bluegill recruitment, growth, population

size structure, and associated factors in Minnesota lakes. North American Journal

of Fisheries Management 25:171-179.

18. Weimer, E.J. 2004. Bluegill seasonal habitat selection, movement, and

relationship to angler locations in a South Dakota glacial lake. M.S. thesis, South

Dakota State University, Brookings.

19. Werner, R.G. 1969. Ecology of limnetic bluegill (Lepomis macrochirus) fry in

Crane Lake, Indiana. The American Midland Naturalist 81:164-181.

20. Willis, D.W., D.A. Isermann, M.J. Hubers, B.A. Johnson, W.H. Miller, T.R.

St.Sauver, J.S. Sorensen, E.G. Unkenholz, and G.A. Wickstrom. 2001. Growth of

South Dakota fishes: A statewide summary with means by region and water type.

South Dakota Department of Game, Fish, and Parks, Special Report 01-05, Pierre.

623

Redear Sunfish, Lepomis microlophus (Günther, 1859)

Etymology and Synonyms: Lepomis = Greek, lepis- meaning “scaled”, and –poma meaning “lid”, meaning scaled operculum; microlophus = “small nape”. Also referred to as “shellcrackers”, referring to the species capability of crushing mollusks with their specialized pharyngeal teeth.

Description: Body deep, laterally compressed with arched dorsal profile. Dorsally golden brown to olive; laterally tan to gray with 5-10 faint, dark olive, diffuse vertical bars depending on the size; ventrally yellow-green to cream; fins without prominent spots; lateral sides of head with small, dark brown to dark orange spots; opercle with a dark black center with a red to burnt-orange, crescent-shaped margin on posterior edge.

Head and snout large, conical. Eye moderately large, amber in color, placed laterally on head. Mouth terminal, small; upper jaw does not extend past anterior edge of eye; lower jaw slightly protrudes past upper jaw. Lips fleshy. Pharyngeal teeth moliform. Gill rakers on first gill arch short, stout, 9-11. Opercle flap short, stiff. Dorsal fin elongate with 9-11 spines and 11-12 rays. Adipose fin absent. Caudal peduncle slightly elongate, thick.

Caudal fin slightly forked with rounded lobes. Anal fin with 3 spines and 10-11 rays.

Pelvic fin insertion slightly posterior to insertion of pectoral fin. Pectoral fin elongate, extending past anterior end of eye when folded forward. Lateral line compete, arched upward with 34-45 ctenoid scales in series. Spawning males with bright yellow breast, light orange coloration on the pectoral and pelvic fins, and a bright amber iris of the eye.

Similar Species: Resembles the Pumpkinseed and Bluegill. Pumpkinseed display 3-5 blue-green wavy lines extending backwards to the gill flap, a short, flexible opercle flap with a black center and white cream margin separated by a single red spot on the 624 posterior edge, and a pectoral fin that only extends to the anterior edge of eye when folded forward. Bluegill display a solid dark blue opercle, without any red spots.

Distribution and Habitat: Native from the Mississippi River in Missouri and Indiana, south to North Carolina and Florida, and west into eastern Texas. Have been introduced outside its native range as a means of sportfishing, and occurs in southwestern South

Dakota throughout the Cheyenne River drainage. Inhabits clear, quiet, warm waters of ponds, marshes, lakes and reservoirs, but may also occur in pools, backwaters, or oxbow areas of low gradient streams and large rivers in areas with little current.11 Often associated with aquatic vegetation or submerged structure, which provide cover and feeding benefits. Adults often in deep waters near the bottom. Although they may be negatively affected by turbidity, Redear Sunfish may be more tolerable of turbidity than bass or bluegills.6,11,12

Reproduction: Reproduction similar to other species of Lepomis. Spawning often beings in May, and can continue through August.9 Adults migrate from deeper waters to shallow areas near shorelines to spawn. Sexual maturity is most likely associated with size rather than age.11 Males defend and excavate saucer-shaped nests with their caudal fins. Nests often constructed in colonies along edges of aquatic vegetation or submerged structure in quiet waters at various depths ranging 4-183 cm (0.13-6.00 ft) in areas often exposed to sun with silt, sand, or mud substrate.11 Males known to court females into the nesting site with a series of grunting sounds, made by the jaw and pharyngeal bones.4 Fractional spawners; females spawn in more than one nest throughout the season. Males provide parental care by defending and fanning the eggs until hatching occurs. Optimal water temperature for incubation and successful fry is 21-24°C (69.8-75.2°F), but has been 625 observed at 32.3°C (90.14°F).11 Fecundity dependent on size of female. A female 151 mm (5.9 in) TL can produce 7,500-13,000 eggs, and a 213 mm (8.39 in) TL female may produce 23,000-25,500 eggs.6,12 Eggs roughly 1.5 mm (0.06 in) in diameter.8 Hatching takes place within 6-10 days depending on water temperature.11 Males continue to defend and watch over fry, who remain within the nest for roughly one week post hatching.7,11

Age and Growth: One of the larger species of sunfish. Capable of growing faster and achieving larger sizes than Bluegill where they occur sympatrically.9,10 Populations in turbid waters often experience slower growth than populations in clear waters.1,6,12

Optimal water temperatures for growth is 24-27°C (75.2-80.6°F).5,11 Growth believed to cease when water temperature falls below 10°C (50°F).11 Adults may reach 380 mm

(14.96 in) TL, but are often shorter.6 Longevity 6 years in its northern ranges, but capable of reaching 11 years in the south.6,9,10

Food and Feeding: Larvae and young fish consume zooplankton, green algae and small aquatic insects such as chironomid larvae.2,11 Adults opportunistic and primarily feed on benthic organisms including snails, clams, and mussels, which they are capable of consuming by crushing with their moliform pharyngeal teeth.3,6 Adults also known to consume aquatic insects such as midge larvae.11 Rarely feed along the waters surface.

Literature Cited:

1. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

2. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville. 626

3. Fisher Huckins, C.J. 1997. Functional linkages among morphology, feeding

performance, diet, and competitive ability in molluscivorous sunfish. Ecology

78:2401-2414.

4. Gerald, J.W. 1971. Sound production during courtship in six species of sunfish

(Centrarchidae). Evolution 25:75-87.

5. Hill, L.G., G.D. Schnell, and J. Pigg. 1975. Thermal acclimation and temperature

selection in sunfishes (Lepomis, Centrarchidae). Southwestern Naturalist 20:177-

184.

6. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

7. Lipinot, A. 1961. The red-ear sunfish. Illinois Wildlife 17:3-4.

8. Meyer, F.A. 1970. Development of some larval centrarchids. Progressive Fish

Culturist 32:130-136.

9. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

10. Sammons, S.M., D.G. Partridge, and M.J. Maceina. 2006. Differences in

population metrics between bluegill and redear sunfish: implications for the

effectiveness of harvest restrictions. North American Journal of Fisheries

Management 26:777-787.

11. Twomey, K.A., G. Gebhart, O.E. Maughan, and P.C. Nelson. 1984. Habitat

suitability index models and instream flow suitability curves: Redear sunfish. U.S.

Fish and Wildlife Service FWS/OBS-82/10.79. 29pp. 627

12. Warren, M.L., Jr. 2009. Centrarchid identification and natural history. Pages 375-

533 in S.J. Cooke and D.P. Phillip, editors. Centrarchid fishes: diversity, biology,

and conservation. Wiley-Blackwell, West Sussex, United Kingdom.

628

Smallmouth Bass, Micropterus dolomieu (Lacepède, 1802)

Etymology and Synonyms: Micropterus = small fin, referring to a torn fin on the type specimen; dolomieu = referring to M. Dolomieu, a French mineralogist.

Description: Body fusiform, slender, slightly laterally compressed, moderately deep.

Dorsally brown or olive green; laterally bronze with dark mottling or faint vertical bars; ventrally cream to white; fins dusky with no distinct markings; 3-4 dark bars present across cheek and opercle; dark spot roughly the size of the pupil on posterior edge of opercle. Head large; scales on cheek smaller than scales on opercle. Snout blunt. Eye large, amber. Mouth slightly oblique; adults with maxillary extending to middle or posterior edge of eye, never further; mandible protrudes past maxillary. Teeth small, present in pads on upper and lower jaws; absent on tongue. Gill rakers 6-8. Dorsal fin with two lobes, moderately joined; anterior fin with 9-10 spines, shortest spine more than half the length of the longest spine; posterior fin with 13-15 rays. Adipose fin absent.

Caudal peduncle thick. Caudal fin slightly forked. Anal fin with 3 spines, 10-11 rays.

Pelvic fin insertion anterior to insertion of dorsal fin; 1 spine and 5 rays. Pectoral fin rounded. Lateral line complete; 68-80 ctenoid scales. Spawning males develop darker coloring and a bright red eye. Juveniles similar to adults with more distinct vertical bars laterally; also with prominent dark caudal spot.

Similar Species: Closely resembles the Largemouth bass. Mandible of Largemouth bass extends to posterior edge or further of eye, and the scales on the cheek and opercle are the same size. Largemouth Bass have slightly joined dorsal fins; anterior fin with 9-10 spines, shortest spine less than half the length of the longest spine; posterior fin with 12- 629

13 rays. Adult Largemouth Bass lack dark mottling and vertical bars laterally, and instead display a dark lateral stripe.

Distribution and Habitat: Widely distributed and stocked throughout the United States outside of its native range due to popularity as a sportfish. Native to the east-central

United States in the St. Lawerence, Great Lakes, Hudson (Red River), and Mississippi

River drainages, ranging from the northeast corner of South Dakota (Minnesota River drainage) to New York, and south to northern Alabama and eastern Oklahoma.7 In the

Dakotas, the species was introduced to the Missouri River reservoirs in the early

1980’s.2,10 A warm and cool water species that establishes well-defined home ranges.3,11

Prefers rocky littoral or shoal habitats in lakes, reservoirs, and riffles or pools within small to medium-size streams with moderate current. Most frequent in clear to slightly turbid waters. Adults often associated in deeper off-shore habitat near submerged structure and vegetation. Juveniles prefer near-shore, shallow habitats with slower velocities and overhanging vegetation.

Reproduction: In lower Lake Oahe, Smallmouth Bass >300 mm (11.8 in) were more likely to be captured in the spring during onshore-offshore movements, likely suggesting spawning migrations.9 Spawning takes place in late spring and early summer within quiet, shallow waters near shore. Males defend and construct nests within rocky or sandy substrate near submerged or overhanging structure with the use of their fins. Fractional spawners. Eggs adhesive, demersal, pale-yellow, roughly 2.0-3.5 mm (0.07-0.14 in) in diameter.3 Fecundity varies with size of the female. Hatching takes place within one week. Males give parental care pre- and post-hatching until juveniles disperse from the nest. 630

Age and Growth: Growth rates highly variable. Length at age varies among population and depends on region and habitat. In Lake Poinsett, South Dakota, age-1 individuals exhibited their greatest growth rates during mid-summer, while primarily feeding on macroinvertebrates.6 Lake and reservoir growth rates often more rapid than streams.3

Average length 305-508 mm (12-20 in) TL. Longevity 6-12 years.

Food and Feeding: Apex, opportunistic predators. Feed on a wide variety of abundant or available prey items.6,8 Undergo an ontogenetic diet shift from larvae to adult. Larval

Smallmouth Bass primarily consume zooplankton and chironomids. Juveniles shift to consume larger aquatic macroinvertebrates, crayfish, and small prey fishes. In eastern

South Dakota lakes, adult Smallmouth Bass consumed invertebrates in the spring, and shifted to a prey fish diet of mainly age-0 Yellow Perch, followed by Black Crappie and

Bluegill in mid and late summer.1,4 Smallmouth bass predation on age-0 Yellow Perch does not appear to limit Yellow Perch recruitment in eastern South Dakota lakes.4 In both eastern South Dakota lakes and Lake Sharpe, adult Smallmouth Bass predation rates on

Walleye occur at a minimal rate that is not likely to detrimentally affect Walleye recruitment.1,5,12,13 Smallmouth Bass size and prey fish availability is thought to determine prey fish consumption.1

Literature Cited:

1. Bacula, T.D. 2009. Smallmouth bass seasonal dynamics in northeastern South

Dakota glacial lakes. Master’s thesis, South Dakota State University, Brookings,

South Dakota.

2. Berry, C.R. Jr., and B. Young. 2004. Fishes of the Missouri National Recreational

River, South Dakota, and Nebraska. Great Plains Research 14:89-114. 631

3. Brewer, S.K., and D.J. Orth. 2014. Smallmouth Bass Micropterus dolomieu

Lacepède, 1802. American Fisheries Society Symposium 82:9-26.

4. Dembkowski, D.J. D.W. Willis, B.G. Blackwell, S.R. Chipps, T.D. Bacula, and

M.R. Wuellner. 2015. Influence of smallmouth bass predation on recruitment of

age-0 yellow perch in South Dakota glacial lakes. North American Journal of

Fisheries Management, 35(4):736-747.

5. Galster, B.J., M.R. Wuellner, and B.D.S. Graeb. 2012. Walleye Sander vitreus

and smallmouth bass Micropterus dolomieu interactions: an historic stable-isotope

analysis approach. Journal of Fish Biology 81:135-147.

6. Gangl, R.S., K.L. Pope, and D.W. Willis. 1997. Seasonal trends in food habits and

growth of smallmouth bass in Lake Poinsett, South Dakota. South Dakota Game,

Fish and Parks, Fisheries Division Special Report 97-5, Pierre.

7. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry Jr. 2007.

Zoogeographic patterns and faunal changes of South Dakota fishes. Western

North American Naturalist 67:161-184.

8. Lott, J.P. 1996. Relationships between smallmouth bass feeding ecology and

population structure and dynamics in lower Lake Oahe, South Dakota. South

Dakota Game, Fish and Parks, Wildlife Division, Completion Report No. 96-3,

Pierre.

9. Lott, J.P. 2000. Smallmouth bass movement, habitat use and electrofishing

susceptibility in lower Lake Oahe, South Dakota. South Dakota Game, Fish and

Parks, Wildlife Division, Annual Report No. 00-15, Pierre. 632

10. Milewski, C.L., and D.W. Willis. 1990. A statewide summary of smallmouth bass

sampling data from South Dakota waters. South Dakota Department of Game,

Fish and Parks, Fisheries Report 90-9, Pierre.

11. Todd, B.L., and C.F. Rabeni. 1989. Movement and habitat use by stream-dwelling

Smallmouth Bass. Transcations of the American Fisheries Society 118(3):229-

242.

12. Wuellner, M.R. 2009. Exploring competitive interactions between Walleye and

Smallmouth Bass in South Dakota Waters. Thesis dissertation, South Dakota

State University, Brookings, South Dakota.

13. Wuellner, M.R., S.R. Chipps, D.W. Willis, and W.E. Adams Jr. 2010. Interactions

between walleyes and smallmouth bass in a Missouri River reservoir with

consideration of the influence of temperature and prey. North American Journal

of Fisheries Management, 30(2):445-463.

633

Largemouth Bass, Micropterus salmoides (Lacepède, 1802)

Etymology and Synonyms: Micropterus = small fin, referring to a torn fin on the type specimen; salmoides = derived from salmo, meaning “trout”, referring to a past common name for the species in the southern states.

Description: Body fusiform, robust, slightly laterally compressed, deep. Dorsally dark olive-green to brown; laterally brown to tan, adults with thick, dark, and irregular (zig- zag) lateral stripe; ventrally cream to white; fins tan with no spots or markings; dark opercle spot roughly the size of the pupil. Head large; scales on cheek and opercle similar in size. Snout blunt. Eye large, golden brown. Mouth slightly oblique, large; adults with maxillary extending to posterior edge or further of eye; mandible thick, protruding well past maxillary. Teeth small, conical; present in pads on upper and lower jaws; absent on tongue. Gill rakers 7-9. Dorsal fin with 2 nearly separate lobes; anterior fin with 9-10 spines, shortest spine less than half the length of longest spine; posterior fin with 12-13 rays. Adipose fin absent. Caudal peduncle thick. Caudal fin slightly forked. Anal fin with

3 spines, 10-12 rays. Pelvic fin insertion anterior to insertion of dorsal fin; 1 spine and 5 rays. Pectoral fin rounded. Lateral line complete; 60-68 ctenoid scales. Spawning males develop a darker coloring. Juveniles similar to adults with prominent vertical blotches making up the lateral stripe.

Similar Species: Closely resembles the Smallmouth Bass. The mandible of Smallmouth

Bass rarely extends past the posterior edge of the eye, and the scales on the cheek are smaller than the scales on the opercle. Smallmouth Bass have moderately joined dorsal fins; anterior fin with 9-10 spines, shortest spine more than half the length of the longest 634 spine; posterior fin with 13-15 rays. Adult Smallmouth Bass lack a dark lateral stripe, and instead display mottling and vertical bars laterally.

Distribution and Habitat: Widely distributed and stocked outside of the native range due to its popularity as a sportfish. Native to the St. Lawrence, Great Lakes, Hudson (Red

River), and Mississippi River drainages, south to the Gulf of Mexico, and into northeastern Mexico. Two subspecies of Largemouth Bass are recognized: the Northern

Largemouth Bass, M. s. salmoides, and the Florida Largemouth Bass, M. s. floridanus.

Northern Largemouth Bass native to the Midwestern United States, and within the

Dakotas are native to the Red River, Minnesota, and Big Sioux River drainages.5 A warm water species that inhabits a variety of aquatic ecosystems throughout both states including small ponds, lakes, and reservoirs, as well as low current areas of creeks and backwaters of medium to large sized rivers. Most frequent in clear to slightly turbid, shallow waters with sand or gravel substrates and aquatic vegetation. The presence of submerged macrophytes (20-40%) is often associated with quality Largemouth Bass populations.3 Rare in waters deeper than 6.1 m (20 ft). Moderately tolerant of turbidity and salinity.

Reproduction: In the Dakotas, recruitment and reproductive success is known to vary within region and population.7,8,11 Natural recruitment and reproductive success in many impoundments within the Dakotas are often negatively impacted by various environmental factors including severe winters and droughts.1,7,12 Thus, the species is periodically stocked as fingerlings. Sexual maturity variable; typically 250-300 mm (9.8-

11.8 in).6 Spawning takes place in spring, but is highly dependent on water temperature.

Males defend and construct 61-69 mm (2-3 ft.) in diameter nests in the substrate with the 635 use of their fins, often in shallow margins with submerged or overhanging structure. A range of courtship behaviors such as nudging and nipping take place as the male attracts a female to the nest. Fractional spawners. Eggs adhesive, demersal, pale-yellow in color, and roughly 1.5-1.7 mm (0.05-0.06 in) in diameter.10 Fecundity varies greatly with the size of the female. Hatching takes place within 7 days. Males give parental care pre and post hatching, and are known not to feed during this time.

Age and Growth: Growth rates highly variable and differ among populations and regions. Increased density of Largemouth Bass, particularly in water bodies <40 ha (99 ac), often leads to reduced growth rates, condition, and size structure within the population.9 The presence of Bluegill as a food source and the length of the growing season affect growth rate in South Dakota ponds.4 Increases rapidly in length within first two years; weight gains typically greater after.2 Average length 254-508 mm (10-20 in).

Females often larger than males. Longevity 15-20 years.

Food and Feeding: Apex, keystone, opportunistic predators in many aquatic ecosystems, where they assist in regulating panfish populations.4 Consumes a wide array of prey items including fish, aquatic insects and crustaceans, small terrestrial mammals, and birds.

Feeding takes place both day and night, often found near aquatic vegetation.

Literature Cited:

1. Beck, R.D. 1986. Growth, survival, and reproductive success of largemouth bass

stocked with selected forage fishes in South Dakota ponds. Master’s thesis, South

Dakota State University, Brookings, South Dakota.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 636

3. Durocher, P.P., W.C. Provine, and J.E. Kraai. 1984. Relationship between

abundance of largemouth bass and submerged vegetation in Texas reservoirs.

North American Journal of Fisheries Management 4(1):84-88.

4. Guy, C.S., and D.W. Willis. 1990. Structural relationships of largemouth bass and

bluegill populations in South Dakota ponds. North American Journal of Fisheries

Management 10:338-343.

5. Hoagstrom, C.W., S.S. Wall, J.G. Kral, B.G. Blackwell, and C.R. Berry, Jr. 2007.

Zoogeographic patterns and faunal change of South Dakota fishes. Western North

American Naturalist 67:161-184.

6. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

7. Kolander, T.D. 1992. Factors limiting overwinter survival of young-of-the-year

largemouth bass in South Dakota. Master’s thesis, South Dakota State University,

Brookings, South Dakota.

8. Lucchesi, D.O., T.R. St. Sauver, B.A. Johnson, K.A. Hoffman, and D.W. Willis.

2004. Region III small impoundments strategic plan, 2004-2009. South Dakota

Department of Game, Fish, and Parks Special Report No. 04-13, Pierre.

9. Paukert, C.P., and D.W. Willis. 2004. Environmental influences on largemouth

bass Micropterus salmoides populations in shallow Nebraska lakes. Fisheries

Management and Ecology 11:345-352.

10. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p. 637

11. Spengler, D.E. 2010. Natural reproductive cycle of northern largemouth bass in

the upper Midwest, with applications to off-season spawning. Master’s thesis,

South Dakota State University, Brookings, South Dakota.

12. Willis, D.W., and C.S. Guy. 1991. Largemouth bass management in South

Dakota: comparison with waters further south and east. Pages 336-342 in J.L.

Cooper and R.H. Hamre, editors. Warmwater Fisheries Symposium I, USDA

Forest Service General Technical Report RM-207.

638

White Crappie, Pomoxis annularis (Rafinesque, 1818)

Etymology and Synonyms: Pomoxis = “sharp opercle”, referring to the sharp, pointed opercle or gill flap; annularis = “annular”, or “with rings”, most likely referring to the rings or bars present on the lateral sides.

Description: Body strongly laterally compressed, deep. Dorsally dark olive-gray to bluish-green; laterally silver to light green with 8-10 dark, vertical, mottled bars (paler in individuals inhabiting turbid water); ventrally silver to white; dorsal, caudal, and anal fins heavily mottled with nearly round alternating dark and light colored spots; dark opercle spot present, slightly larger than pupil. Head small, short, strongly concave in profile.

Snout short, pointed. Eye moderately large, placed laterally on head; golden yellow to olive green in color. Mouth large, moderately oblique; upper jaw extending at least to middle of eye; lower jaw extending past upper jaw. Teeth present on both jaws in pads; small, conical. Gill rakers long, slender, 25-32. Dorsal fin large, rounded with 5-6 spines and 12-16 rays; base slightly shorter than or equal to base of anal fin. Adipose fin absent.

Caudal peduncle slender, slightly elongate. Caudal fin slightly forked. Anal fin large, rounded with 5-6 spines and 16-18 rays. Pelvic fin insertion well anterior to insertion of dorsal fin. Pectoral fin with 14-16 rays. Lateral line complete with 36-46 ctenoid scales.

Spawning males generally with more pronounced, dark coloring. Juveniles similar to adults with a more elongate and less deep body.

Similar Species: Closely resembles the Black Crappie. Black Crappie display heavier, dark mottling on lateral sides, with vertical bars absent, as well as a dorsal fin with 7-8 spines and 14-16 rays. Black Crappie also lack a prominent dark opercle spot. 639

Distribution and Habitat: Native to the extreme eastern edge of South Dakota and

Minnesota in the north, east to New York, and south to the Gulf of Mexico. Have been widely introduced in the United States outside its native range due to popularity as a sportfish. Less common than Black Crappie in the Dakotas. Found most often schooling in warm waters such as lakes, ponds, and reservoirs, but also occur in low gradient creeks, and moderate to large-sized streams. Optimum water temperature 27-28.5°C

(80.6-83.3°F).1 In lentic environments, White Crappie are often found at mid-depths during the day, and retreat to shallow waters near shore during the evening and night.5,7

Monthly and diel movement patterns likely related to feeding behavior, and may vary with habitat and lake morphology.5 Home ranges in a South Dakota glacial lake varied from <0.1-85.0 ha (0.25-210.04 a).5 In lotic environments, the species frequents areas with low velocity such as pools and backwaters. Prefer to inhabit areas with submerged cover and structure or aquatic vegetation with sand, mud, gravel, or silt substrates. More tolerant of turbidity and siltation than Black Crappie.7,9 Overwinter in deeper depths, where they will remain until spring.

Reproduction: Reproductive behavior similar to Black Crappie. Spawning takes place earlier in southern populations, and spans from May to June in the Dakotas, or when water temperature reaches 14-23°C (57-73°F).1 Spawning sites often in shallow waters with shelter or vegetation. Sexual maturity is reached at age 2-3. Males excavate littoral nests or small depressions arranged in colonies in depths of 0.2-6.1m (7.87-240.16 in).7,10

Aggressive and territorial behavior is exhibited by males when defending the nests.1

Males provide parental care by guarding and fanning the nest until larvae hatch, and fry leave the nest within a couple days. Fecundity increases with age and condition of 640 female.2 Females produce roughly 10,000-180,000 eggs per season.11 Eggs roughly 0.89 mm (0.03 in) in diameter, demersal, and adhesive to the substrate or aquatic vegetation around the nest.1,10 Hatching occurs within 2-5 days depending on water temperature.10

Age and Growth: Growth rates highly variable among populations. During the first years of life, growth rates are density dependent and stunting may occur when the species is overabundant.8 Mean length-at-age from multiple South Dakota waters were reported as: age-1, 93 mm (3.66 in) TL; age-2, 183 mm (7.20 in) TL; age-3, 221 mm (8.70 in) TL; age-4, 252 mm (9.92 in) TL; age-5, 275 mm (10.83 in) TL.13 Capable of reaching up to

381-508 mm (15-20 in) TL. Longevity 3-12 years, with few surviving past age-5.

Food and Feeding: Feeding behavior and diet similar to Black Crappie. Young mainly feed during daylight hours on suspended zooplankton such as rotifers, cladocerans and copepod nauplii.1,6 Majority of adult feeding takes place between dusk and dawn when individuals migrate from deeper to shallower waters. Adults primarily feed near or just below the surface, or among aquatic vegetation. Adult diets consist of larger zooplankton such as Daphnia, insects such as Chironomids and Corixids, and small fish such as age-0

Gizzard Shad.3,12 Fish are added to the diet when individuals reach roughly 150 mm (5.91 in) TL.4,6 Small fish are more important to the diet of White Crappie than the Black

Crappie.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Bunnell, D.B., M.A. Scantland, and R.A. Stein. 2005. Testing for evidence of

maternal effects among individuals and populations of white crappie.

Transactions of the American Fisheries Society 134:607-619. 641

3. Busiahn, T.R. 1977. Food, growth, and reproduction of white crappies (Pomoxis

annularis) and black crappies (P. nigromaculatus) in Lake Poinsett, South

Dakota. M.S. Thesis, South Dakota State University, Brookings.

4. Ellison, D.E. 1984. Trophic dynamics of a Nebraska black crappie and white

crappie population. North American Journal of Fisheries Management 4:355-364.

5. Guy, C.S., and D.W. Willis. 1994. Biotelemetry of white crappies in a South

Dakota glacial lake. Transactions of the American Fisheries Society 123:63-70.

6. O’Brien, W.J., B. Loveless, and D. Wright. 1984. Feeding ecology of young

white crappie in a Kansas reservoir. North American Journal of Fisheries

Management 4:341-349.

7. Pflieger, W.L. 1975. The fishes of Missouri. Missouri Department of

Conservation. 343 p.

8. Pope, K.L., G.R. Wilde, and B.W. Durham. 2004. Age-specific growth patterns in

density-dependent growth of white crappie, Pomoxis annularis. Fisheries

Management and Ecology 11:33-38.

9. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

10. Siefert, R.E. 1968. Reproductive behavior, incubation and mortality of eggs, and

postlarval food selection in the white crappie. Transactions of the American

Fisheries Society 97:252-259.

11. Smith, P.W. 1979a. The fishes of Illinois. University of Illinois Press, Urbana. 642

12. Unkenholz, D. 1971. Food habits of black crappies, white crappies, yellow perch,

and white suckers in a small impoundment in northeastern South Dakota. M.S.

Thesis, South Dakota State University, Brookings.

13. Willis, D.W., D.A. Isermann, M.J. Hubers, B.A. Johnson, W.H. Miller, T.R.

St.Sauver, J.S. Sorensen, E.G. Unkenholz, and G.A. Wickstrom. 2001. Growth of

South Dakota fishes: A statewide summary with means by region and water type.

South Dakota Department of Game, Fish, and Parks, Special Report 01-05, Pierre.

643

Black Crappie, Pomoxis nigromaculatus (Lesueur, 1829)

Etymology and Synonyms: Pomoxis = “sharp opercle”, referring to the sharp, pointed opercle or gill flap; nigromaculatis = nigro, meaning “black”, and maculates, meaning

“spotted”, referring to the species coloration.

Description: Body strongly laterally compressed, deep. Dorsally dark olive to bluish- green; laterally light olive to silvery green with heavy, irregular dark green and silver speckled mottling (paler in individuals inhabiting turbid water); ventrally silver to white; dorsal, caudal, and anal fins heavily mottled with nearly round alternating dark and light colored spots. Head small, short, strongly concave in profile. Snout short, pointed. Eye moderately large, placed laterally on head; yellow-brown in color. Mouth large, terminal and oblique; upper jaw extending at least to middle of eye; lower jaw extending past upper jaw. Teeth present on both jaws in pads; small, conical. Gill rakers long, slender,

27-32. Dorsal fin large, rounded with 7-8 spines and 14-16 rays; base roughly equal to base on anal fin. Adipose fin absent. Caudal peduncle slender, slightly elongate. Caudal fin slightly forked. Anal fin large, rounded with 6-7 spines and 17-18 rays. Pelvic fin insertion well anterior to insertion of dorsal fin. Pectoral fin with 13-15 rays. Lateral line complete with 38-44 ctenoid scales. Spawning males generally with more pronounced, dark coloring. Juveniles similar to adults with a more elongate and less deep body.

Similar Species: Closely resembles the White Crappie. White Crappie display a dorsal fin with 5-6 spines and 12-16 rays, and have lighter lateral sides with 8-10 dark, vertical, mottled bars present. White Crappie also display a dark opercle spot.

Distribution and Habitat: Native from Manitoba, Canada in the north, south to

Minnesota continuing to the Gulf of Mexico, and east to the Atlantic coast. Not assumed 644 to be native to North or South Dakota, but have been widely introduced in the United

States outside the native range due to popularity as a sportfish. Often found schooling in warm waters such as lakes, ponds, and reservoirs, as well as low gradient creeks, streams, and rivers. In lentic environments, the species frequents mid-depths during the day, and migrates to shallow waters near shore during the evening and night.5 In lotic environments, Black Crappie are often found in pools, backwaters or areas with low velocity. Prefer clear water with abundant cover or aquatic vegetation. Less tolerant of turbidity and siltation than White Crappie.11 Migrations to deeper depths and overwintering habitat begin in the late fall, where they will remain until spring.

Reproduction: Spawning takes place earlier in southern populations, and spans from

May to June in the Dakotas, or when water temperature reaches 14.4-20°C (58-68°F).1

Spawning sites often in sheltered, vegetated bays of lakes. Sexual maturity is reached at age 2-3. Males excavate nests or small depressions in colonies, roughly 200-230 mm

(7.87-9.05 in) in diameter in depths of 0.4-0.8 m (15.75-31.50 in) with gravel, silt, or sand substrate, in areas that are protected from wind and wave action, usually by some sort of structure like woody debris and cattails.1,9 Males provide parental care by guarding and fanning the nest until larvae hatch, and fry leave the nest within a couple days. Males remain in nesting sites for several weeks.9 Fecundity increases with age; mean fecundity from females age 3-4 in Lake Poinsett, South Dakota was 46,364 eggs per female.2 Eggs roughly 0.93 mm (0.04 in) in diameter and adhesive to the substrate or aquatic vegetation around the nest.6 Hatching occurs within 2-4 days.7

Age and Growth: Growth highly variable among ecosystems.3 Turbidity known to reduce growth and survival of juveniles.8 Populations with low density known to grow 645 faster than populations with high density.2,4 In South Dakota, growth known to be slower in smaller than in large impoundments, and fastest growth rates occur in natural lakes.3

Mean length-at-age from multiple South Dakota waters are reported as: age-1, 77 mm

(3.03 in) TL; age-2, 151 mm (5.94 in) TL; age-3, 198 mm (7.79 in) TL; age-4, 227 mm

(8.94 in) TL; age-5, 248 mm (9.76 in) TL; age-6, 262 mm (10.31 in) TL; age-7, 264 mm

(10.39 in) TL.3 Capable of reaching up to 381 mm (15 in) TL. Longevity 6-9 years.

Food and Feeding: Feed by sight, and prey upon the largest, slowest moving organisms requiring the least bit of energy.2 Majority of feeding takes place in the morning and night when individuals migrate from deeper to shallower waters. Small larval Black

Crappie <10 mm (0.39 in) TL and medium sized larvae 10-29 mm (0.39-1.14 in) TL select calanoid copepods; medium and large sized larvae 10-59 mm (0.39-2.32 in) TL prey on Daphnia.10 Prey selectivity of larval Black Crappie in addition to zooplankton availability and vulnerability may determine year-class strength.10 Adults mainly feed near or just below the surface, or among aquatic vegetation, and mainly feed on zooplankton such as Daphnia, insects such as Corixids, and small fish.2 Small fish are less important to the diet of Black Crappie than the White Crappie.

Literature Cited:

1. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

2. Busiahn, T.R. 1977. Food, growth, and reproduction of white crappies (Pomoxis

annularis) and black crappies (P. nigromaculatus) in Lake Poinsett, South

Dakota. M.S. Thesis, South Dakota State University, Brookings. 646

3. Guy, C.S., and D.W. Willis. 1993. Statewide summary of sampling data for black

and white crappies collected from South Dakota waters. South Dakota

Department of Game, Fish, and Parks, Completion Report 93-12, Pierre.

4. Guy, C.S., and D.W. Willis. 1995. Population characteristics of Black Crappies in

South Dakota waters: A case for ecosystem-specific management. North

American Journal of Fisheries Management 15:754-765.

5. Guy, C.S., R.M. Neumann, and D.W. Willis. 1993. Seasonal and diel movements

of adult black crappies in a South Dakota natural lake. South Dakota Department

of Game, Fish, and Parks, Completion Report 93-14, Pierre.

6. Merriner, J.V. 1971. Egg size as a factor in intergeneric hybrid success of

centrarchids. Transactions of the American Fisheries Society 100:29-32.

7. Pflieger, W.L. 1975. The fishes of Missouri. Missouri Department of

Conservation. 343 p.

8. Pope, K.L. 1996. Factors affecting recruitment of black crappies in South Dakota

waters. Doctoral dissertation, South Dakota State University, Brookings.

9. Pope, K.L., and D.W. Willis. 1997. Environmental characteristics of black crappie

(Pomoxis nigromaculatus) nesting sites in two South Dakota waters. Ecology of

Freshwater Fish 6:183-189.

10. Pope, K.L., and D.W. Willis. 1998. Early life history and recruitment of black

crappie (Pomoxis nigromaculatus) in two South Dakota waters. Ecology of

Freshwater Fish 7:56-68.

11. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p. 647

CHAPTER 22

FAMILY PERCIDAE

Introduction

The Perch family, Percidae, consists of 10 genera and more than 200 species native to North America and Eurasia. In the Dakotas, this family is famously recognized by the popular and important game-fish species, Walleye (Sander vitreus), Yellow Perch

(Perca flavescens), and Sauger (Sander Canadensis); however, the majority of species in this family are much smaller, and are known collectively as darters. As with the general pattern of fish diversity, there is a greater number of darter species in the southern portion of North America, specifically east of the Rocky Mountains. Five species of darters occur in the Dakotas. The family Percidae also includes other non-game species such as the

Logperch (Percina caprodes), which is also found in both eastern North and South

Dakota. Although native to western Eurasia, Zander (Sander lucioperca) were introduced into Spiritwood Lake, North Dakota in 1989 for sport fishing. Although they are not plentiful, they have established a population within the lake, and yearlings to age-2 individuals are still being recorded today.

Members of the family Percidae may sometimes be confused with members of the

Temperate Bass family (Moronidae) and the Sunfish family (Centrarchidae). Percids can easily be differentiated from the two families by having one or two anal spines, whereas

Temperate Bass possess three anal spines, and Sunfish have three or more anal spines.

Furthermore, Percids can also be distinguished from Temperate Bass by the absence of well-developed opercle spines, and a lateral line that does not extend onto the upper lobe 648 of the caudal fin. Percids are also recognized by having an anterior spinous dorsal fin, and a posterior rayed dorsal fin, which are often times distinctly separated from each other. Even though they belong to the same family, the larger Percid species like

Walleye, Yellow Perch and Sauger are not often confused with darters, even in juvenile form, as they possess easily recognizable teeth on both sets of jaws. Darter have much smaller teeth that are not as easily seen. The absence of a swim bladder is also an internal characteristic that separates darters from the larger Percids, which have fully developed and functional swim bladders. The lack of a swim bladder allows the darters to be bottom-dwelling fishes, often perched upon the bottom substrate, unlike the larger

Percids, which inhabit the pelagic zone.

Percids in the Dakotas are found within a wide variety of habitat types. The larger

Percids primarily inhabit lakes, reservoirs, and rivers, while the majority of the darter species prefer creeks, streams and lakes. Many larger Percid species undergo an ontogenetic diet change throughout their lifetime, primarily consuming invertebrates as juveniles before switching to a primarily piscivorous diet as adults. Darters on the other hand, generally rely on aquatic invertebrates throughout the entirety of their lifetime.

Spawning patterns of Percids also differentiate between the larger species and darters.

Walleye, Yellow Perch and Sauger mainly exhibit random or broadcast spawning behavior over gravel or rocky substrate where they do not construct a nest or provide parental care. Darters are more specific when it comes to choosing a place to spawn, and most all darter species provide some form on parental care to either their eggs, fry, or both.

649

Iowa Darter, Etheostoma exile (Girard, 1859)

Etymology and Synonyms: Etheostoma: Greek, etho = to strain + Greek, stoma = mouth

(a straining mouth); exile: Latin = slim.

Description: Body elongate, slender, slightly laterally compressed. Sexually dimorphic species. Adult males olive brown to dark brown on dorsal side fading to yellowish white on ventral side; roughly 8 dark blotches along dorsal side; 9 – 12 prominent vertical bars on lateral sides, with dark red to burnt orange blotches in interspaces; distinct black suborbital bar; spiny dorsal fin with three color bands: basal band slate gray, middle band broad and light colored, and outer band dusky brown; soft dorsal fin light in color with small dark spots arranged in rows; caudal fin with small dark spots giving a wavy appearance; anal, pelvic and pectoral fins transparent in color with dark pigment along rays. Head moderate. Snout blunt. Eye large. Mouth small, slightly oblique to horizontal.

Premaxillae nonprotractile. Upper lip groove non continuous over snout. Frenum present.

Teeth tiny, sharp, placed in narrow bands on upper and lower jaws. Infraorbital canal with 8 pores. Gill rakers short, roughly 7. Dorsal fins 2: anterior spiny dorsal fin with 8 –

12 spines; soft posterior dorsal fin with 10 – 12 rays. Caudal peduncle long. Caudal fin squarish with rounded corners. Anal fin with 2 spines and 7 – 9 rays. Pelvic fin base anterior to dorsal fin insertion. Pectoral fin fan shaped. Lateral line incomplete with 55 –

65 scales in the series; fewer than 35 pored. Scales ctenoid; opercles, nape and cheeks scaled; belly scaled; breast slightly scaled. Adult female much like adult male in appearance, but without bright colors; lateral bars less distinct; often times more olive to yellow-brown. Breeding males display 9 – 12 bright blue lateral bars with red interspaces; spiny dorsal fin layered with bright blue spots along basal half, then bright 650 orange to red band, with narrow blue stripe on outer edge. No tubercles displayed on breeding males.

Similar Species: The Iowa Darter differs from the Johnny Darter (Ethestoma exile),

Blackside Darter (Percina maculata), River Darter (Percina shumardi), and Slenderhead

Darter (Percina phoxocephala) by lacking a complete lateral line. Johnny Darter display numerous small X’s and W’s along the lateral sides. Blackside Darter have connected black blotches along the lateral line creating a stripe. River Darter lack a frenum, or is absent when mouth is closed. Slenderhead Darter display a pointed snout.

Distribution and Habitat: Distributed throughout south central Canada, and in the north central United States from Montana east to the Great Lakes, south to the Hudson Bay

River drainages. Reported in the Sheyenne and Souris rivers, Fort Peck, Lake Francis

Case, and smaller streams of the Missouri and Little Missouri River drainages in North

Dakota.5 In South Dakota, it occurs in all principle drainages east of the Missouri River, and in the Niobrara River drainage to the west.1 Resides in calm or standing semi turbid to clear waters, such as natural lakes and ponds, as well as slow moving streams. Often associated with heavy amounts of submerged aquatic vegetation and filamentous algae covered surfaces.2 Found over sand, gravel, and silty substrates.

Reproduction: Migration from deeper portions of lakes and streams to shallower shorelines takes place in late spring. Migration can be postponed due to colder weather and water temperatures. Males migrate to spawning grounds prior to females. Spawning takes place over fibrous root material and rooted aquatic vegetation near shorelines from late April to June.2,5 Males establish spawning territories, in the shape of a semicircle with the base along the shoreline.2 Defensive behavior is exhibited by males only towards 651 their own species when defending their territories, often by erecting their fins.2 When the female is ready to spawn, a single male courts her to the spawning site where he mounts her anteriorly with his posterior end curved downward around the female. Their bodies vibrate while eggs are fertilized and deposited. Often times, less than 10 eggs are deposited at a single spawning episode, with roughly 900 total eggs being deposited throughout the spawning season.3 Females spawn with several males during the season, and return to deeper water when finished.2 Males continue to defend the spawning territory after the eggs have been deposited.

Age and Growth: Females often larger than males. Adults average 45.72 – 63.5mm (1.8

– 2.5 in.) TL, sometimes reaching 69.85mm (2.75in.) Can live up to 4 years. Length and age in northern Wisconsin was documented as: age 1, 34 – 55mm (1.33 – 2.17in.); age 2,

48 – 61mm (1.88 – 2.40in.); age 3, 58 –68mm (2.28 – 2.67in.); age 4, 68 – 69mm (2.67 –

2.72in.) TL.2,4

Food and Feeding: Adults feed primarily on aquatic insects and larvae, mayflies, chironomids, dipterans, and sometimes fish eggs. Juveniles feed on small insects and crustaceans, including copepods and amphipods.

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL. 652

4. Lutterbie, G.W. 1976. The darters (Pices: Percidae: Etheostomatinae) of

Wisconsin. University of Wisconsin Stevens Point. MAT Thesis. 307pp.

5. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

653

Johnny Darter, Etheostoma nigrum (Rafinesque, 1820)

Etymology and Synonyms: Etheostoma = Greek, etho-, meaning “to strain”, -stoma, meaning “mouth” (a straining mouth); nigrum = Latin, “black”, possibly referring to the dark coloring of spawning males.

Description: Body fusiform, elongate, slender, slightly laterally compressed. Dorsally brown to golden-tan with 4-7 dark saddles; laterally light brown to tan with dark “W” or

“X” shaped markings; ventrally creamy yellow to white; dorsal and caudal fin spotted, other fins lightly pigmented to transparent. Head moderate. Snout blunt. Eye large, placed dorsolaterally on head. Opercle partially to completely scaled. Mouth terminal to slightly subterminal; upper jaw extends to anterior end of eye. Frenum absent. Teeth small, sharp, present in bands on upper and lower jaws. Gill rakers short, stout, 5-11. Dorsal fin with two lobes scarcely joined; anterior lobe with 8-10 spines, posterior lobe with 11-14 rays.

Adipose fin absent. Caudal peduncle elongate, slender. Caudal fin square to slightly rounded. Anal fin with 1 spine and 7-9 rays. Pelvic fins thoracic. Pectoral fin with 10-14 rays. Lateral line complete with 35-50 ctenoid scales in series. Spawning males develop dusky gray to black on head, anal, and pelvic fins; pelvic fin spine and rays develop white tips. Juveniles similar to adults.

Similar Species: Easily distinguished from other darter species in the Dakotas by the dark “W”, and “X” lateral markings. Iowa Darter lack a complete lateral line and display a distinct black suborbital bar. Blackside Darter display 6-8 large, irregular, oval shaped blotches on lateral sides to the base of the caudal fin. River Darter lack a frenum and display a distinct black suborbital bar. Slenderhead Darter have a pointed snout and an anal fin with 2 spines. 654

Distribution and Habitat: Native throughout east-central North America from

Colorado, southeastern Montana, and Saskatchewan in the west, east throughout the

Missouri, Mississippi, Great Lakes, and Hudson Bay drainages to Quebec and Virginia, and south throughout the Atlantic Slope and Mobile Bay drainages to Alabama and

Mississippi. Occurs throughout the major tributaries of the Missouri River in North

Dakota, and the major tributaries east of the Missouri River in South Dakota. Inhabits quiet, shallow areas such as pools or moderately flowing runs of streams, rivers, and lakes with moderate aquatic vegetation and sand or gravel substrate. Intolerant of high turbidity levels, and tolerant of low dissolved oxygen and high water temperatures.7,11

Critical thermal maximum roughly 31°C (87.8°F).6

Reproduction: Spawning takes place April to June, or when water temperatures reach

11.7-21.1°C (53.06-69.98°F).2 Spawning may take place earlier in southern latitudes.10

Quick changes in temperature, increased siltation, and rising water levels may postpone or interrupt spawning.2 Fractional spawners. Males migrate to spawning areas prior to females and establish nests by brushing off sediment with their fins from the underside of submerged structures, such as rocks.1,14,15 Nests located in areas with slow current, often in pools or shallow waters of lakes.1,14 Either sex may initiate spawning when in close proximity by turning themselves upside down to attract one another.1 Spawning act consists of both male and female in an inverted position against a rock or the submerged structure covering the nest, quivering in a side-by-side or head-to-tail position.1,14

Females rarely extract more than one egg at a time.14 Nests contain 30-1,589 eggs.4,14

Fecundity increases with size and age of female, with females depositing 50-200 eggs per season.2,13 Eggs amber with dark oil globule, roughly 1.5 mm (0.06 in) in diameter.1,13 655

Males provide parental care until hatching by fanning, rubbing, and defending the eggs from predators.1,5 Darting motions, an open mouth, and quivering are all behaviors of males chasing predators from the nest.1 Eggs covered in fungus often consumed by the male. Hatching occurs within 6 days at 23°C (73.4°F), 10 days at 20°C (68°F), and 16 days at 13°C (55.4°F).13

Age and Growth: Newly hatched larvae roughly 5 mm (0.20 in) TL.2 Growth rapid during first two years. Mean lengths-at-age from Iowa are reported as: age-1, 28.8 mm

(1.13 in) TL; age-2, 43 mm (1.69 in) TL; age-3, 53 mm (2.09 in) TL; age-4, 56 mm (2.20 in) TL.8 Males typically larger than females. Average length 38.1-63.5 mm (1.5-2.5 in)

TL, but capable of reaching 77 mm (3.03 in) TL.2 Longevity 4 years.8

Food and Feeding: Visual predator adapted to forage on sedentary benthic prey.9,12

Majority of feeding takes place during the day.3 A protrusible premaxillae allows the upper jaw to extend well beyond the lower jaw to help consume prey.9 Adults primarily feed on chironomid larvae and aquatic insects such as diptera, ephemeroptera, trichoptera, and odonata.8,9 Also known to consume microcrustaceans such as ostracods, cladocerans, and copepods, but to a lesser extent.8,9

Literature Cited:

1. Atz, J.W. 1940. Reproductive behavior in the eastern Johnny darter, Boleosoma

nigrum olmstedi (Storer). Copeia 2:100-106.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Emery, A.R. 1973. Preliminary comparisons of day and night habits of freshwater

fish in Ontario lakes. Journal of the Fisheries Research Board of Canada 30:761-

774. 656

4. Grant, J.W.A., and P.W. Colgan. 1983. Reproductive success and mate choice in

the Johnny darter, Etheostoma nigrum (Pisces: Percidae). Canadian Journal of

Zoology 61:437-446.

5. Grant, J.W.A., and P.W. Colgan. 1984. Territorial behavior of the male Johnny

darter, Etheostoma nigrum. Environmental Biology of Fishes 10:261-269.

6. Ingersoll, C.G., and D.L. Claussen. 1984. Temperature selection and critical

thermal maxima of the fantail darter, Etheostoma flabellare, and Johnny darter, E.

nigrum, related to habitat and season. Environmental Biology of Fishes 11:131-

138.

7. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

8. Karr, J.R. 1963. Age, growth, and food habits of Johnny, slenderhead and

blacksided darters of Boone County, Iowa. Proceedings of the Iowa Academy of

Science 70:228-236.

9. Paine, M.D., J.J. Dodson, and G. power. 1981. Habitat and food resource

partitioning among four species of daters (Percidae: Etheostoma) in a southern

Ontario stream. Canadian Journal of Zoology 60:1635-1641.

10. Parrish, J.D., D.C. Heines, and J.A. Baker. 1991. Reproductive season, clutch

parameters, and oocyte size of the Johnny darter Etheostoma nigrum from

southwestern Mississippi. The American Midland Naturalist 125:180-186.

11. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City. 657

12. Roberts, N.J., and H.E. Winn. 1962. Utilization of the senses in feeding behavior

of the Johnny darter, Etheostoma nigrum. Copeia 1962:567-570.

13. Speare, E.P. 1965. Fecundity and egg survival of the central Johnny Darter

(Etheostoma nigrum nigrum) in southern Michigan. Copeia 3:308-314.

14. Winn, H.E. 1958a. Observation on the reproductive habits of darters (Pisces-

Percidae). American Midland Naturalist 59:190-212.

15. Winn, H.E. 1958b. Comparative reproductive behavior and ecology of fourteen

species of darters (Pisces-Percidae). Ecological Monographs 28:155-191.

658

Yellow Perch, Perca flavescens (Mitchill, 1814)

Etymology and Synonyms: Perca = Greek for “a perch”; flavescens = “yellowish”, referring to the species coloration on lateral sides.

Description: Body fusiform, laterally compressed, moderately deep, oval in cross section. Dorsally brown to olive with 6-9 dark, saddles that extend down vertically on lateral sides; laterally yellow-green to golden bronze; ventrally white; dorsal fin with dusky gray to black pigment on first 2 and last 3-5 membranes; anal, pelvic, and pectoral fins pale yellow to light orange-red. Head small, elongate. Snout conical, blunt. Eye with gold iris, placed laterally on head. Mouth moderately large, terminal, slightly oblique; upper jaw extends to pupil of eye; jaws nearly equal in length, with lower jaw occasionally slightly longer than upper jaw. Frenum absent. Teeth small, pointed, present in pads on both jaws. Gill rakers 18-20. Dorsal fin with two distinctly separate lobes; anterior fin with 12-14 spines; posterior dorsal fin with 1-2 spines, and 12-15 rays.

Adipose fin absent. Caudal peduncle slender, elongate. Caudal fin moderately forked.

Anal fin with 2 spines, and 7-8 rays. Pelvic fins thoracic, insertion posterior to insertion of posterior dorsal fin. Pectoral fins with 13-15 rays. Lateral line complete with 52-60 ctenoid scales. Spawning adults similar to non-spawners. Juveniles similar to adults with less deep bodies.

Similar Species: No other species within the Dakotas closely resemble the Yellow Perch.

Sunfish much deeper bodied, with two lobes of dorsal fin distinctly joined. Bass lack saddles, or vertical bars on lateral sides, and have an anal fin with 3 spines. Walleye and

Sauger possess large, distinct canine teeth. 659

Distribution and Habitat: Native to central Canada and the United States east of the

Rocky Mountains from North Dakota and east-central South Dakota in the west, east throughout the Mississippi, Great Lakes, and Atlantic River basins, and south to South

Carolina, Ohio, and Nebraska. Adaptable to a variety of habitat types, the species occurs widely throughout the Dakotas, and are often found schooling in clear to slightly turbid waters of lakes, ponds, sloughs, reservoirs, and streams with submerged aquatic vegetation and/or submerged structure over sand and gravel substrates. Eurythermal, temperate species, with an optimal water temperature ranging between that of a cold- and warmwater fish, primarily at 20-27°C (68-80.6°F).3,23

Reproduction: Spawning takes place early spring, with greatest egg viability at 8-11°C

(46.4-51.8°F).13 Water temperatures later in the spring are most optimal for spawning and larval development than earlier in the spring.22 Preferred spawning habitat in an eastern

South Dakota lake consists of areas within the main body of the lake, approximately 0.6 m (1.97 ft) in depth within 3-30 m (9.84-98.4 ft) from shore, with rocky and gravel substrate and periphyton-free submerged wood structure.7 Sexual maturity occurs earlier in populations with good growth, and is generally reached at age-2 for males and age-3 for females.11,19,32 Spawning activity mainly takes place at night and occurs over a 5-11 day period.1,17 No nest is prepared and no parental care is given. Females release a gelatinous, tubular strand of eggs, which become entangled within submerged vegetation and structure.7 Fecundity highly variable, and increases with size and age of female.12

Females >350 mm (13.7 in) TL are capable of producing 100,000-150,000 eggs.19 Eggs

1.0-2.1 mm (0.04-0.08 in) in diameter.34 Warmer water temperatures increase hatching success and create a shorter hatching period with fewer abnormalities in the larvae.14 660

Hatching occurred at 9 days at 13.3°C (55.94°F), 23-24 days at 8.7-10.0°C (47.66-50°F), and 35 days at 7.7°C (45.86°F).15 Recruitment often inconsistent and highly variable among water bodies and is characterized by strong, weak, or missing year-classes.18,25,27

Annual recruitment of larvae is primarily driven by abiotic factors including lake morphology, fluctuations in water levels, and especially temperature.4,5,16,22,30

Recruitment known to increase in years with increased precipitation and less variation in temperatures.22,30 Biotic factors such as prey availability, prey size are also known to influence annual recruitment.6,20

Age and Growth: Larvae 5-6 mm (0.20-0.24 in) TL at time of hatching.34 Optimal growth for age-0 individuals occurs at 23°C (73.4°F).6 Thermal requirements vary among populations geographically, but in South Dakota, maximum growth for length and weight occurs in water temperature 24.8-25.4°C (76.6-77.7°F).2,3 Growth of Yellow Perch has been frequently correlated with abiotic factors such as temperature and fluctuation in water levels, however it is mostly influenced by biotic factors such as prey availability, predation, and intra- and intrespecific competition.4,5,21,22,28,29,31,33 Statewide mean lengths-at-age from South Dakota are reported as: age-1, 86 mm (3.39 in) TL; age-2, 145 mm (5.71 in) TL; age-3, 190 mm (7.48 in) TL; age-4, 220 mm (8.66 in) TL; age-5, 242 mm (9.53 in) TL.35 Capable of reaching 406.4 mm (16 in) TL and 0.91 kg (2 lbs).

Longevity 10 years.

Food and Feeding: Generalist predators. Yellow Perch larvae primarily feed on zooplankton such as rotifers and copepod nauplii until roughly 7.0 mm (0.28 in) TL.6

Small larvae 20-40 mm (0.79-1.57 in) TL mainly consume adult copepods and cladocerans.10 Intermediate larvae 40-60 mm (1.57-2.36 in) TL, positively select benthic 661 macroinvertebrates, which provide a higher energetic gain.6,10 Survival and growth of larvae may increase with periods of high copepod densities.9 Adult diets in South Dakota are highly variable among location and month, however chironomids are a major source of prey in all populations.26 Prey fish are introduced in the diet around age-1, or at 80-150 mm (3.15-9 in) TL, and individuals are not primarily piscivorous until age-4.8,10,24

Literature Cited:

1. Breder, C.M., Jr., and D.E. Rosen. 1966. Modes of reproduction in fishes.

American Museum of Natural History, New York.

2. Brown, M.L., and K.A. Smith. 2004. Temperature-dependent growth models for

South Dakota yellow perch, Perca flavescens, fingerling production. Journal of

Applied Aquaculture 16:105-112.

3. Brown, P.B., J.E. Wetzel, J. Mays, K.A. Wilson, C.S. Kasper, and J. Malison.

2002. Growth differences among stocks of yellow perch, Perca flavescens, are

temperature dependent. Journal of Applied Aquaculture 12:43-56.

4. Dembkowski, D.J., M.J. Weber, and M.R. Wuellner. 2017. Factors influencing

recruitment and growth of age-0 yellow perch in eastern South Dakota glacial

lakes. Fisheries Management and Ecology 24:372-381.

5. Dembkowski, D.J., S.R. Chipps, and B.G. Blackwell. 2014. Response of walleye

and yellow perch to water-level fluctuations in glacial lakes. Fisheries

Management and Ecology 21:89-95.

6. Fisher, S.J., and D.W. Willis. 1997. Early life history of yellow perch in two

South Dakota glacial lakes. Journal of Freshwater Ecology 12:421-429. 662

7. Fisher, S.J., K.L. Pope, L.J. Templeton, and D.W. Willis. 1996. Yellow perch

spawning habitats in Pickerel Lake, South Dakota. Prairie Naturalist 28:65-75.

8. Fullhart, H.G., B.G. Parsons, D.W. Willis, and J.R. Reed. 2002. Yellow perch

piscivory and its possible role in structuring littoral zone fish communities in

small Minnesota lakes. Journal of Freshwater Ecology 17:37-43.

9. Graeb, B.D.S., J.M. Dettmers, D.H. Wahl, and C.E. Cáceres. 2004. Fish size and

prey availability affect growth, survival, prey selection, and foraging behavior of

larval Yellow Perch. Transactions of the American Fisheries Society 133:504-

514.

10. Graeb, B.D.S., M.T. Mangan, J.C. Jolley, D.H. Wahl, and J.M. Dettmers. 2006.

Ontogenetic changes in prey preference and foraging ability of yellow perch:

insights based on relative energetic return of prey. Transactions of the American

Fisheries Society 135:1493-1498.

11. Hasler, A.D. 1945. Observations on the winter perch population of Lake Mendota.

Ecology 26:90-94.

12. Heyer, C.J., T.J. Miller, F.P. Binkowski, E.M. Caldarone, and J.A. Rice. 2001.

Maternal effects as a recruitment mechanism in Lake Michigan yellow perch

(Perca flavescens). Canadian Journal of Fisheries and Aquatic Sciences 58:1477-

1487.

13. Hokanson, K.E.F. 1977. Temperature requirements of some percids and

adaptations to the seasonal temperature cycle. Journal of the Fisheries Research

Board of Canada 34:1524-1550. 663

14. Hokanson, K.E.F., and C.F. Kleiner. 1974. Effects of contstant and rising

temperatures on survival and developmental rates of embryonic and larval yellow

perch, Perca flavescens (Mitchill). Pages 437-448 In: Blaxter, J.H.S. (ed.), The

early life history of fishes. Springer-Verlag, New York.

15. Huff, D.D., G. Grad, and C.E. Williamson. 2004. Environmental constraints on

spawning depth of yellow perch: the roles of low temperature and high solar

ultraviolet radiation. Transactions of the American Fisheries Society 133:718-726.

16. Isermann, D.A. 2003. Population dynamics and management of yellow perch

populations in South Dakota glacial lakes. Doctoral dissertation, South Dakota

State University, Brookings.

17. Isermann, D.A., and D.W. Willis. 2008. Emergence of larval Yellow Perch, Perca

flavescens, in South Dakota lakes: potential implications for recruitment.

Fisheries Management and Ecology 15:259-271.

18. Isermann, D.A., D.W. Willis, B.G. Blackwell, and D.O. Lucchesi. 2007. Yellow

perch in South Dakota: population variability and predicted effects of creel limit

reductions and minimum length limits. North American Journal of Fisheries

Management 27:918-931.

19. Jansen, W.A. 1996. Plasticity in maturity and fecundity of yellow perch, Perca

flavescens (Mitchill): comparisons of stunted and normal-growing populations.

Annales Zoologici Fennici 33:403-415.

20. Jolley, J.C., D.W. Willis, and R.S. Holland. 2010. Match-mismatch regulation for

bluegill and yellow perch larvae and their prey in Sandhill lakes. Journal of Fish

and Wildlife Management 1:73-85. 664

21. Kaemingk, M.A., and D.W. Willis. 2012. Mensurative approach to examine

potential interactions between age-0 yellow perch (Perca flavescens) and bluegill

(Lepomis macrochirus). Aquatic Ecology 46:353-362.

22. Kaemingk, M.A., B.D.S. Graeb, and D.W. Willis. 2014. Temperature, hatch date,

and prey availability influence age-0 yellow perch growth and survival.

Transactions of the American Fisheries Society 143:845-855.

23. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

24. Keast, A. 1985. The piscivore feeding guild in small freshwater ecosystems.

Environmental Biology of Fishes 12:119-129.

25. Koonce, J.F., T.B. Bagenal, R.F. Carline, K.E.F. Hokanson, and M. Nagiec. 1977.

Factors influencing year-class strength of percids: a summary and a model of

temperature effects. Journal of the Fisheries Research Board of Canada 34:1890-

1899.

26. Lott, J.P. 1991. Food habits of yellow perch in eastern South Dakota glacial lakes.

M.S. thesis, South Dakota State University, Brookings.

27. Newsome, G.E., and S.K. Alto. 1987. An egg-mass census method for tracking

fluctuations in yellow perch (Perca flavescens) populations. Canadian Journal of

Fisheries and Aquatic Sciences 44:1221-1232.

28. Paukert, C.P., D.W. Willis, and J.A. Klammer. 2002. Effects of predation and

environment on quality of yellow perch and bluegill populations in Nebraska

Sandhill lakes. North American Journal of Fisheries Management 22:86-95. 665

29. Pierce, R.B., C.M. Tomcko, and M.T. Negus. 2006. Interactions between stocked

walleyes and native yellow perch in Lake Thirteen, Minnesota: a case history of

percid community dynamics. North American Journal of Fisheries Management

26:97-107.

30. Pope, K.L., D.W. Willis, and D.O. Lucchesi. 1996. Differential relations of age-0

black crappie and yellow perch to climatological variables in a natural lake.

Journal of Freshwater Ecology 11:345-350.

31. Power, M., and M.R. van den Heuvel. 1999. Age-0 yellow perch growth and its

relationship to temperature. Transactions of the American Fisheries Society

128:687-700.

32. Purchase, C.F., N.C. Collins, G.E. Morgan, and B.J. Shuter. 2005. Sex-specific

covariation among life-history traits of yellow perch (Perca flavescens).

Evolutionary Ecology Research 7:549-566.

33. Schoenebeck, C.W., and M.L. Brown. 2010. Potential importance of predation,

competition, and prey on yellow perch growth from two dissimilar population

types. The Prairie Naturalist 42:32-37.

34. Thorpe, J.E. 1977. Morphology, physiology, behavior, and ecology of Perca

fluviatilis L. and Perca flavescens Mitchill. Journal of the Fisheries Research

Board of Canada 34:1504-1514.

35. Willis, D.W., D.A. Isermann, M.J. Hubers, B.A. Johnson, W.H. Miller, T.R. St.

Sauver, J.S. Sorensen, E.G. Unkenholz, and G.A. Wickstrom. 2001. Growth of

South Dakota fishes: A statewide summary with means by region and water type.

South Dakota Department of Game, Fish, and Parks, Special Report 01-05, Pierre.

666

Logperch, Percina caprodes (Rafinesque, 1818)

Etymology and Synonyms: Percina = Latin, a diminutive of Perca, meaning “Perch”; caprodes = Greek for “pig-like”, in reference to the shape of the species snout.

Description: Body fusiform, elongate, slightly laterally compressed. Dorsally olive to tan; laterally tan to light olive with 14-25 dark olive to gray vertical bars of alternating long and short lengths; ventrally cream to white; suborbital present, may be dark to faint; dorsal and caudal fins with dark, dusky speckles forming bands; dark spot present at base of caudal fin. Head moderate. Cheeks, opercles, and nape of neck scaled. Snout pointed, conical, elongate, overhangs mouth. Eye moderately large, placed laterally on upper portion of head. Mouth subterminal, horizontal; upper jaw does not extend to anterior edge of eye. Frenum present. Lips fleshy. Teeth small, sharp, present on both jaws. Gill rakers short, 12-15. Anterior dorsal fin with 13-15 spines; posterior dorsal fin with 14-17 rays. Adipose fin absent. Caudal peduncle elongate, thick. Caudal fin square to slightly forked. Anal fin with 2 spines, and 9-11 rays. Pelvic fins thoracic. Pectoral fins with 13-

15 rays. Lateral line complete with 76-82 ctenoid scales in series. Sexually dimorphic species. Males with enlarged anal fin and single row of enlarged modified scales ventrally along the midline between pelvic fin insertion and vent. Females typically lack enlarged modified scales ventrally. Spawning males with minute tubercles present on posterior ventral side of body, caudal peduncle, caudal, anal, pelvic and pectoral fins.

Juveniles similar to adults.

Similar Species: May be confused with the Blackside and Slenderhead Darter. Blackside

Darter have a shorter, less dramatic pointed snout, and display 6-8 somewhat large and irregular oval shaped blotches, almost connected on lateral sides. Slenderhead Darter lack 667 the distinct 14-25 dark olive to gray vertical bars of alternating lengths that Logperch display, and instead are distinguished by a light yellow-tangerine to orange submarginal band on the spinous dorsal fin.

Distribution and Habitat: Native to much of east central North America from

Saskatchewan and the Upper Mississippi River drainage in the west, east through the

Great Lakes and St. Lawrence river drainage to Quebec and Vermont in the east, and south throughout the Mississippi River systems to Louisiana and the Rio Grande River in southern Texas.3 Native to the Red, Minnesota, and Big Sioux River drainages of eastern

North and South Dakota. Inhabit lakes within eastern South Dakota, but are also known to occur in medium to large sized streams and rivers in riffle areas with gravel substrate.

1,16 Often found schooling within a variety of habitat types including areas with little to swift current such as pools, riffles and sandy shoals, and prefers clear to slightly turbid water.16 Frequently occurs over sand, gravel, and cobble substrates in depths 0.6-1.5 m

(1.97-4.92 ft).2,7 Have been known to occupy depths of 9-40 m (29.52-131.23 ft) in the

Great Lakes.2,13,15

Reproduction: Spawning takes place early morning to early evening in April-early

July.2,3,4,5,6 Sexual maturity reached at age 1-2.2 Ripe females enter spawning grounds, swimming through schools of males when ready to spawn.3 In lakes, spawning grounds consist of areas up to 30 m (98.43 ft) in length parallel to the shoreline with sand or gravel substrate.16 Spawning grounds much smaller in streams and rivers.16 One or more males initiate the spawning act by courting a single female and create a “moving territory”.16 The male clasps the female with the pelvic fins in an upright horizontal position with their tails pressed together, and their bodies vibrate to initiate the release 668 and fertilization of eggs.3,16 This behavior often causes the pair and eggs to become almost completely buried in the bottom substrate.16 No nest is prepared, and no parental care is given, leaving exposed eggs to be frequently consumed by other male

Logperch.2,16 Fractional spawners; females spawn with more than one male during the season.3 After spawning ceases, adults migrate back to deeper waters.16 Fecundity increases with size and age of the female, and ranges 1,000-3,000.16 Eggs adhesive, demersal, amber in color with several small and one large oil globule, and roughly 1.12 mm (0.04 in) in diameter.3 Hatching occurred within 200-205 hours after fertilization when eggs reached 1.27 mm (0.05 in) in diameter in an aquarium with an average water temperature of 16.5°C (61.7°F).3

Age and Growth: Newly hatched larvae roughly 4.5-5.3 mm (0.18-0.21 in) TL.11 Larvae classified as juveniles at 18-24 mm (0.71-0.94 in) TL.3,11 Average lengths at age from

Illinois are reported as: age-1, 71.5 mm (2.81 in) TL; age-2, 105.9 mm (4.17 in) TL; age-

3, 121.5 mm (4.78 in) TL.12 Capable of reaching 180 mm (7.09 in) TL.13 Longevity 4 years.

Food and Feeding: Sedentary, littoral, benthic, generalized omnivore.8,14 Little seasonal variation in food habits.10 Adults mainly consume aquatic insect larvae, pupae and adults, fish eggs, amphipods, small crustaceans, and small mollusks.2,10,16 Forages by using its conical snout to turn over pebbles and rocks.9 Larvae and young begin as surface feeders, mainly consuming microcrustacea such as copepods and cladocerans.14

Literature Cited:

1. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor. 669

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Cooper, J.E. 1978. Eggs and larvae of the Logperch, Percina caprodes

(Rafinesque). The American Midland Naturalist 99:257-269.

4. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

5. Eddy, S., and J.C. Underhill. 1974. Northern fishes with special reference to the

upper Mississippi Valley. Third edition. University of Minnesota Press,

Minneapolis

6. Lutterbie, G.W. 1976. The darters (Pisces: Percidae: Ethestomatinae) of

Wisconsin. M.S. Thesis, University of Wisconsin, Stevens Point.

7. Morris, M.A., and L.M. Page. 1981. Variation in western logperches (Pisces:

Percidae), with description of a new subspecies from the Ozarks. Copeia 1981:95-

108.

8. Mullan, J.W., R.L. Applegate, and W.C. Rainwater. 1968. Food of logperch

(Percina caprodes), and Brook Silverside (Labidesthes sicculus), in a New and

Old Ozark Reservoir. Transactions of the American Fisheries Society 97:300-305.

9. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

10. Phillips, E.C., and R.V. Kilambi. 1996. Food habits of four benthic fish species

(Ethestoma spectabile, Percina caprodes, Noturus exilis, Cottus carolinae) from

northwest Arkansas streams. The Southwestern Naturalist 41:69-73. 670

11. Simon, T.P. 1985. Descriptions of larval Percidae inhabiting the upper Mississippi

River Basin (Osteichthys: Ethestomatini). M.S. Thesis, University of Wisconsin,

Lacrosse.

12. Thomas, D.L. 1970. An ecological study of four darter species of the genus

Percina (Percidae) in the Kaskaskia River, Illinois. Illinois Natural History

Survey, Biological Notes 70:1-18.

13. Trautman, M.B. 1957. The fishes of Ohio. Ohio State University Press,

Columbus. 638pp.

14. Turner, C.L. 1921. Food of the common Ohio darters. Ohio Journal of Science

22:41-62.

15. Wells, L. 1968. Seasonal depth distribution of fish in southeastern Lake

Michigan. U.S. Fish and Wildlife Service Fisheries Bulletin 67:1-15.

16. Winn, H.E. 1958. Comparative reproductive behavior and ecology of fourteen

species of darters (Pisces-Percidae). Ecological Monographs 28:155-191.

671

Blackside Darter, Percina maculate (Girard, 1859)

Etymology and Synonyms: Percina = Latin for Perca meaning perch; maculate = spotted, derived from the Latin word maculates, referring to the lateral blotches.

Description: Body elongate, moderately robust and slightly laterally compressed. Dorsal side olive green to light brown and yellowish in color with wavy checkerboard design and 6-11 dusky crossbars; ctenoid scales; lateral sides yellow with lateral stripe black to brown running from the snout, through the eye, leading into 6-8 somewhat large and irregular oval shaped blotches, almost connected, running longitudinal to the body to the base of the caudal fin; blotches more connected in young than adults. Suborbital bar vertical and dusky. Ventral side yellowish white. Head pointed. Snout short and pointed.

Opercle and cheeks with embedded scales; nape and anterior portion of ventral side with few to none embedded scales. Mouth terminal to subterminal and slightly oblique; upper jaw extending into below front of eye; upper lip groove non-continuous over point of snout; teeth small and placed in bands along upper and lower jaws. Frenum present, wide on upper lip. Gill rakers on first arch roughly ten; long and narrow. Dorsal fins 2; anterior spiny dorsal fin with dark pigment along the anterior base fading posteriorly with 12-15 spines and a dark dusty spot on first three membranes; posterior dorsal fin with 12-15 rays. Caudal fin squarish with light bar pattern; dark spot at caudal base more apparent in young. Anal fin clear to yellowish white with 2 spines and 9-11 rays. Pelvic fins clear to yellowish in color with 1 larger, modified scale at base; males with a thicker and sturdier spine on anterior edge of pelvic fin than females. Pectoral fins light in color. Lateral line complete with 62-68 scales. Small air bladder present internally. Breeding males with darker fins; dark brown iris known to turn gold in color; tubercles absent. 672

Similar Species: The Slenderhead Darter (Percina phoxocephala) displays an orange band in the anterior dorsal fin. The Johnny Darter (Etheostoma nigrum) shows numerous large and obvious “X” and “W” markings along the lateral sides of the body. The Iowa

Darter (Etheostoma exile) displays an incomplete lateral line, unlike the Blackside Darter which has a complete lateral line. The River Darter (Percina shumardi) lacks a frenum, or is not visible when the mouth is closed; lateral line also has 46-62 scales.

Distribution and Habitat: The Blackside Darters range is widespread across the middle eastern portion of North America. It spans from southern Ontario and Manitoba down into the Mississippi River basin from Minnesota to Louisiana, and east to Arkansas and

Alabama. It is found throughout the Great Lakes and Hudson Bay regions, as well as the

Mobile basin and Tennessee River drainage.1 The range spans into the very eastern parts of North Dakota and South Dakota. An established population is known to live in the Big

Sioux River drainage in South Dakota.3,6 There have also been reports of the species occurring in Whetstone Creek, located in the northeastern part of South Dakota flowing into Minnesota waters, the lower James and Souris Rivers, the Sheyenne River and tributaries of the Red River in North Dakota.2,4,5 The Blackside Darter prefers to reside near undercut banks in shallow pools, raceways and riffles in medium to large sized streams with slow to moderate flow.1,3,7 It is rare in lakes. It is often found in clear to slightly turbid water with vegetation present and sand or gravel substrate. Unlike most darters which tend to swim and rest along the substrate, the Blackside Darter utilizes its small swim bladder to swim in the middle of the water column during the day, and then rest on the bottom during night.8,9 Juveniles prefer to hide within vegetation to limit chances of predation. 673

Reproduction: Blackside Darters reach sexual maturity at roughly two years of age.

Very rarely are yearling fish mature.7 The species is known to migrate up tributaries in late spring to spawn.7 Spawning generally spans from April to June and takes place in depressions with gravel or sand substrate in slow currents in roughly one to two feet of water.7,10 Females initiate spawning by swimming nervously into a depression in the sand or gravel substrate. Males create a ‘moving’ territory around the targeted female and are defensive to other males of the same species. A male will then follow the female and clasp onto her. The pair will vibrate their bodies together in a horizontal position for several seconds to release eggs and sperm.7 Fertilized eggs are adhesive, roughly 2mm

(0.08in) in diameter, and clear or transparent in color with a colorless oil globule.7 Eggs are deposited down into the depression where they remain unattended until they hatch.7,10

In general, fecundity is known to range from 1,000-2,000 eggs per female within the spawning season; females spawn with several males throughout the season laying at least

10 eggs each time.7 Egg size varies little with the number of eggs in the ovary and the size of the adult.7 Eggs are known to hatch in roughly 6-8 days, and larvae will drift within the water column for up to three weeks.3

Age and Growth: Newly hatched Blackside Darter measure approximately 5.75mm

(0.22in) TL.10 Age 1 individuals in Iowa measured 35mm (1.4in) to 47-51mm (1.9-2.0in) in Wisconsin.9,11,12 Age 4 individuals in Iowa measured 77-79mm (3.0-3.1in), and 101-

108mm (4.0-4.3in) in Wisconsin.9,11,12 Generally larger than other darters. Males and females are known to be very similar in TL, varying from 50.8-109.22mm (2-4.3in) TL and average size 71mm (2.8in).4,10,11 Blackside Darters can live up to 4 years.11,12 674

Food and Feeding: Blackside Darters mainly consume their prey on or just below the surface of the water and consume mayfly, caddisfly and blackfly larvae, chironomids, copepods, and small amounts of plant material.4,9,10 They have also been known to eat other small fish. Juveniles primarily consume very small crustaceans such as copepods.

Literature Cited:

1. Mettee, M.F., P.E. O’Neil, and J.M. Pierson. 1996. Fishes of Alabama and the

Mobile Basin. Geological Survey of Alabama Monograph 15, Tuscaloosa.

2. Churchill, E.P., and W.H. Over. 1938. Fishes of South Dakota. Brown & Saenger

Printers, Sioux Falls.

3. Hrabik, R.A., S.C. Schainost, R.H. Stasiak, and E.J. Peters. 2015. The Fishes of

Nebraska. Conservation and Survey Division, School of Natural Resources, UNL.

4. Owen, J.B., D.S. Elsen, and G.W. Russell. 1981. Distribution of Fishes in North

and South Dakota Basins affected by the Garrison Diversion Unit. University

Press of University of North Dakota, Grand Forks.

5. Copes, F.A. 1965. Fishes of the Red River tributaries of North Dakota. M.S.

Thesis, University of North Dakota. 65p.

6. Bailey, R.M., and M.O. Allum. 1962. Fishes of South Dakota. University of

Michigan, Museum of Zoology, Ann Arbor.

7. Winn, H.E. 1958b. Comparative reproductive behavior and ecology of fourteen

species of darters (Pices-Percidae). Ecological Monographs 28: 155-191.

8. Gilbert, C.R., and J.D. Williams. 2002. National Audubon Society Field Guide to

Fishes, Revised Edition, North America. Alfred A. Knopf, Inc., New York. 675

9. Kansas Fishes Committee. 2014. Kansas Fishes. University Press of Kansas,

Lawrence.

10. Petravicz, W.P. 1938. The breeding habits of the Black-sided Darter, Hadropterus

maculatus Girard. Copeia 1938 (1): 40-44.

11. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

12. Karr, J.R. 1963. Age, growth, and food habits of johnny, slenderhead, and

blackside darters of Boone County, Iowa. Proceedings of the Iowa Academy of

Science 70: 228-36.

676

Slenderhead Darter, Percina phoxocephala (Nelson, 1876)

Etymology and Synonyms: Percina = Latin, a diminutive of Perca, meaning “Perch”; phoxocephala = “tapered head”, referring to the conical head and pointed snout.

Description: Body elongate, slightly laterally compressed. Dorsally light brown to tan; laterally light olive to tan with 10-16 faint, dark, dusky blotches; ventrally cream to white; dark, backward extending lateral stripe on head from tip of snout through eye; light yellow-tangerine to orange submarginal band on spinous dorsal fin; dark caudal spot present at base of caudal fin. Head elongate. Opercle and nape of neck scaled. Snout conical, moderately pointed. Eye large, placed dorsolaterally on head. Mouth terminal to slightly subterminal; upper jaw extends to anterior end of eye. Frenum present. Teeth small, sharp, present in bands on upper and lower jaws. Gill rakers 9-15. Dorsal fin with

11-2 spines, followed by 11-14 rays. Adipose fin absent. Caudal peduncle elongate, slender. Caudal fin square to slightly forked. Anal fin with 2 spines and 8-9 rays. Pelvic fins thoracic. Pectoral fin with 12-15 rays. Lateral line complete with 60-75 ctenoid scales in series. Sexually dimorphic species. Male with scaled breast on posterior end only, with enlarged scales present from the mid-pelvic area to the anus. Females with naked breast except for 1 or 2 large ctenoid scales. Spawning males with bright orange submarginal band on spinous dorsal fin. Juveniles similar to adults, but may have a larger head and eyes in relation to body size.

Similar Species: Easily distinguished from other darter species in the Dakotas by the light yellow-tangerine to orange sub marginal band on the spinous dorsal fin. Iowa Darter lack a complete lateral line and display a distinct black suborbital bar. Blackside Darter display 6-8 large, irregular, oval shaped blotches on lateral sides to the base of the caudal 677 fin. River Darter lack a frenum and display a distinct black suborbital bar. Johnny Darter display dark “W” or “X” shaped markings on lateral sides and have a more blunt snout.

Distribution and Habitat: Native to North America from the Minnesota River drainage in northeastern South Dakota in the west, east to throughout the Upper Mississippi River drainage to western Pennsylvania, although mainly absent from the Great Lakes drainage, and south to northern Alabama and northern Oklahoma.4 Rare in South Dakota and absent from North Dakota. Most common in areas with sand or gravel substrate in shallow, clear to turbid waters near riffles or swift current in small to medium-sized rivers and streams of moderate gradient.1,4,7 Migrate to deeper areas of the channel post spawning and remain throughout winter months.4 Intolerant of high siltation levels, and more tolerant of turbidity than other darter species.3,6,7

Reproduction: Spawning takes place April to June, or when water temperature hovers

21.1°C (70°F) over riffles with gravel substrate in depths 15-60 cm (0.49-1.97 ft) deep.4

High water levels and varying changes in seasonal temperature may postpone or interrupt spawning.4,5 Males migrate to spawning locations prior to females in late March-early

April, and likely establish territories.4 Spawning colors develop more quickly in larger males.4 Sexual mature individuals range 40-80 mm (1.57-3.15 in) SL.4 Fecundity 50-

1,000 eggs per female, with larger females producing more eggs.2,4 Eggs transparent with one large oil droplet, roughly 1.3 mm (0.05 in) in diameter, and adhesive.4 Hatching occurs within 14 days. Juveniles remain in spawning habitat for roughly one month post hatching before migrating to deeper depths of the main channel.4

Age and Growth: Growth rapid early in life. Larvae roughly 22 mm (0.87 in) TL at two weeks of age.4 Mean lengths at age from Iowa are reported as: age-1, 34.1 mm (1.34 in) 678

TL; age-2, 46.6 mm (1.83 in) TL; age-3, 49.4 mm (1.94 in) TL; 56.0 mm (2.20 in) TL.2

Capable of reaching approximately 100 mm (3.94 in) TL.7 Little relation between size and sex.4 Longevity 4 years.

Food and Feeding: Primarily feeds throughout the day, with little feeding occurring at night.5 Majority of annual feeding activity takes place prior to spawning, and the least amount of feeding takes place after spawning and during winter months.4 Juveniles and young primarily feed in midge larvae.5 Adults consume a variety of prey including midge, black fly, caddisfly and mayfly larvae.4 Also known to consume terrestrial insects, fish eggs, and amphipods, but less frequently.4

Literature Cited:

1. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

2. Karr, J.R. 1963. Age, growth, and food habits of Johnny, slenderhead and

blacksided darters of Boone County, Iowa. Proceedings of the Iowa Academy of

Science 70:228-236.

3. Miller, R.J., and H.W. Robinson. 2004. Fishes of Oklahoma, revised edition.

University of Oklahoma Press, Norman.

4. Page, L.M., and P.W. Smith. 1971. The life history of the slenderhead darter,

Percina phoxocephala, in the Embarras River, Illinois.

5. Thomas, D.L. 1970. An ecological study of four darter species of the genus

Percina (Percidae) in the Kaskaskia River, Illinois. Illinois Natural History

Survey, Biological Notes 70:1-18. 679

6. Thompson, B.A. 1980. Percina phoxocephala (Nelson), slenderhead darter. Page

737 in D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and

J.R. Stauffer Jr., editors. Atlas of North American freshwater fishes. North

Carolina State Museum of Natural History, Raleigh.

7. Trautman, M.B. 1957. The Fishes of Ohio. Ohio State University Press. 683p.

680

River Darter, Percina shumardi (Girard, 1859)

Etymology and Synonyms: Percina = Latin, a diminutive of Perca, meaning “Perch”; shumardi = referring to George C. Shumard, a surgeon and naturalist who worked for the

U.S. Pacific Railroad Survey and discovered the species.

Description: Body fusiform, elongate, slightly laterally compressed. Dorsally dark olive- brown with 5-7 dark diffuse saddles or large irregular shaped spots; laterally light olive to tan with 8-13 dark lateral bars anteriorly and forming blotches posteriorly; ventrally cream to white; anterior dorsal fin with distinct, dark gray to black spot on first membrane posterior to first spine, as well as a dark blotch present on membranes on the posterior end; posterior dorsal and caudal fins with small speckles forming bars; caudal spot present, faint; thick, black suborbital bar present. Head elongate. Opercle scaled.

Nape of neck with little to no scales. Snout short, rounded. Eye moderately large, placed dorsolaterally on head. Mouth terminal to slightly subterminal, slightly oblique; upper jaw extends to anterior edge of eye. Frenum absent or weakly visible when mouth is closed. Teeth small, sharp, present in bands on upper and lower jaws. Gill rakers short,

12-17. Dorsal fin with 10-12 spines, followed by 11-16 rays. Adipose fin absent. Caudal peduncle elongate, slender. Caudal fin square to slightly forked. Anal fin with 2 spines and 10-13 rays. Pelvic fins thoracic. Pectoral fins with 13-15 rays. Lateral line complete with 46-60 ctenoid scales in series. Sexually dimorphic species. Males with enlarged anal fin and modified ventral scales between pelvic fins and vent. Females lack or have fewer ventral scales. Spawning males generally darker in color with small tubercles present on head, pelvic, anal, and caudal fins. 681

Similar Species: Easily distinguished from other darter species in the Dakotas by the thick, black suborbital bar and distinct dark gray to black spot and blotch on anterior dorsal fin. Blackside Darter also display a dark, dusky suborbital bar, however they have a more pointed head and display 6-8 large and irregular oval shaped blotches running longitudinal on the lateral sides. Iowa Darter also have a distinct suborbital bar, but display 9-12 prominent vertical bars on lateral sides and dorsal fins with bands of color.

Johnny Darter display dark “W” or “X” shaped markings on lateral sides. Slenderhead

Darter have a light yellow-tangerine to orange sub marginal band on the spinous dorsal fin and have a conical, moderately pointed snout.

Distribution and Habitat: Native to North America from Manitoba and the Red River basin of North Dakota in the northwest, east to Ontario, the Mississippi River, Great

Lakes, and Hudson Bay drainages and Ohio, south through the Mobile Bay River basin to the Gulf of Mexico. Not known in South Dakota. Inhabit both lentic and lotic habitats, specifically large rivers, lakes, and reservoirs. Smaller streams also provide suitable habitat, especially during winter months and the spawning season.4 Occur most frequently in areas with moderate to swift current such as chutes and riffles, 2-5 m (6.56-

16.40 ft) in depth over sand, gravel, or cobble substrates.7,8,11 Also inhabits rocky shorelines of lakes and reservoirs.2,4 River Darters from Manitoba and Ontario were sampled in water temperatures 8.5-15.6°C (47.3-60.08°F), and have also been historically observed in temperatures up to 25.9°C (78.62°F).7 Juveniles inhabit areas with little to moderate current over sand or gravel substrate alongside the main channel.9 Tolerant of continuously high turbidity levels.6,7 682

Reproduction: Spawning in North Dakota likely takes place April-June. The reproductive season in Manitoba was reported as June-July, April-June in Wisconsin,

April-May in Illinois, and April in Kansas.2,3,8,11 Spawning behavior initiated at a water temperature of 10°C (50°F) in the upper Mississippi River basin.9 Sexual maturity reached at age-1.7,11 Information regarding the spawning act is scarce, however spawning occurs over gravel or rubble substrate in areas with high velocity at depths >0.5 m (1.64 ft).9 Fractional spawner, depositing more than one clutch of eggs during a season.9 Eggs translucent with single oil globule, roughly 1.2 mm (0.05 in) in diameter, demersal, slightly adhesive, and typically buried within sand and gravel.2,9 Hatching in an aquarium was observed within 6-7 days at a water temperature of 22°C (71.6°F).9

Age and Growth: Newly hatched larvae approximately 3.6-4.0 mm (0.14-0.16 in) TL.9

Young-of-year individuals from Illinois ranged 36-43 mm (1.42-1.69 in) in June, and mean lengths-at-age from southern Wisconsin were reported as: age-1, 50 mm (1.97 in), and age-2, 67 mm (2.64 in).5,11 Individuals from Manitoba and northwestern Ontario exhibited slow growth, growing roughly 10 mm (0.39 in) per year.7 Average length 58-84 mm (2.28-3.31 in) TL; capable of reaching 89 mm (3.5 in) TL.6 Longevity 4 years.7,10

Food and Feeding: Feeding takes place during daylight hours, and diet may change seasonally depending on availability.7 Adults mainly consume Diptera, Trichoptera,

Ephemeroptera, Crustacea, fish eggs, and zooplankton.1,7,11 Juveniles and young primarily consume microcrustaceans. Snails have also been noted as an important component of the diet.1,7

Literature Cited: 683

1. Balesic, H. 1971. Comparative ecology of four species of darters (Etheostominae)

in Dauphin and its tributary, the Valley River. M.S. Thesis, University of

Manitoba, Winnipeg.

2. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

3. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

4. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

5. Lutterbie, G.W. 1976. The darters (Pisces: Percidae; Ethestomatinae) of

Wisconsin. M.S. Thesis, University of Wisconsin, Stevens Point.

6. Pflieger, W.L. 1997. The fishes of Missouri, revised edition. Missouri Department

of Conservation, Jefferson City.

7. Pratt, T.C., W.M. Gardner, D.A. Watkinson, and L.D. Bouvier. 2016. Ecology of

the river darter in Canadian waters: distribution, relative abundance, life-history

traits, diet, and habitat characteristics. Diversity 8(4):22.

8. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

9. Simon, T.P. 1985. Descriptions of the larval Percidae inhabiting the upper

Mississippi River basin (Osteichthyes: Etheostomatini). M.S. thesis, University of

Wisconsin, Lacrosse.

10. Smith, P.W. 1979. The fishes of Ohio. University of Illinois Press, Urbana,

Illinois. 684

11. Thomas, D.L. 1970. An ecological study of four darter species of the genus

Percina (Percidae) in the Kaskaskia River, Illinois. Illinois Natural History

Survey, Biological Notes 70:1-18.

685

Sauger, Sander canadensis (Griffith & Smith, 1834)

Etymology and Synonyms: Sander = “pike-perch”, referring to the genus’s resemblance to the non-related pike family (Esocidae); canadensis = “of Canada”, referring to the species native distribution.

Description: Body fusiform, elongate, cylindrical in cross section but slightly laterally compressed. Dorsally dark gray to brown with dark, diffuse saddles extending down lateral sides; laterally dark gray to golden brown with small, speckled dark spots; ventrally cream to white; anterior dorsal fin with distinct, black spots arranged in 3 horizontal rows; posterior dorsal fin with rows of small, dark spots; eye glassy, silver, representing a reflecting layer (tapetum lucidum). Head large, elongate. Snout conical, blunt. Cheek scaled. Eye large, placed laterally on head. Mouth large, terminal, oblique; upper jaw extends to posterior edge of eye; jaws nearly equal in length. Teeth large, sharp, canine, present on both jaws. Gill rakers 7-11. Dorsal fin with two distinctly separate fins; anterior fin with 12-13 spines; posterior fin with 1-2 spines and 17-19 rays.

Adipose fin absent. Caudal peduncle elongate, moderately thick. Caudal fin slightly forked. Anal fin with 2 spines and 11-13 rays; insertion posterior to insertion of posterior dorsal fin. Pelvic fins thoracic. Pectoral fins with 14-16 fin rays. Lateral line compete with 77-87 ctenoid scales. Spawning males similar to non-spawning adults. Juveniles more elongate.

Similar Species: Closely resemble and occur sympatrically with Walleye. Walleye are less elongate with a slightly deeper body, have a cheek with little to no scales, lack a spotted anterior dorsal fin, and have a white tip on the lower lobe of the caudal fin. 686

Distribution and Habitat: Native to North America from southeastern Alberta,

Montana, and Wyoming in the west, to southwestern Quebec and Vermont in the northeast, and south to the southern Mississippi River drainage in Mississippi, Louisiana, and the Tennessee River in northern Alabama.2,10 Populations have declined throughout much of their range due to hybridization, exploitation, loss of spawning areas, and general habitat alterations like construction of dams, which block access to historical spawning sites and alter thermal and hydrologic regimes.1,8,11,12,13 Populations within the reservoirs of the Dakotas however are rather stable.1 Cool-water species, with an optimal water temperature of 19.6-22°C (67.28-71.6°F).13 Young-of-year inhabit areas <3.66 m

(12 ft) in depth.5 Primarily a large river species occurring in deep, highly turbid waters within the main channel over sand and silt substrates, but also inhabit large, shallow lakes.13,14

Reproduction: Highly migratory species; capable of migrating up to 260 km (161.56 mi) upstream in the middle Missouri River.13 Preferred spawning habitat in the Missouri

River mainstem reservoirs consists of areas <1.5 m (4.92 ft) in depth with warmer water temperatures, flowing water and high turbidity in delta habitat and secondary channels or tributary streams.1 Sexual maturity occurs at age 3-4, with females maturing later than males.7 Spawn at temperatures similar to or warmer than walleyes.13 Spawning in Lewis and Clark Lake, South Dakota takes place from late April to early May, and is initiated when water temperature reaches 6.11°C (47°F).5 Spawning takes place during the evening and peak spawning activity last for 5-7 days, but may continue for 14 days.5

Adults retreat back to original location after the spawning act ceases. No nests are constructed, and no parental care is given. Broadcast spawners. Fecundity increases with 687 increasing fish length, however egg quality decreases.16 Females from Lewis and Clark

Lake averaged 65,250 eggs/kg (29,659 eggs/lb) of body weight.7 Eggs roughly 1.0-1.9 mm (0.04-0.07 in) in diameter, demersal, and adhesive before water hardening.5,18

Hatching in Lewis and Clark Lake took place at 21 days at an average water temperature of 8.33-8.70°C (46.94-47.66°F).4,15 In South Dakota Missouri River reservoirs, annual recruitment may be higher in years with warmer spring and early summer water temperatures and reduced flow.1 Year-class strength dependent on fluctuating water levels.5

Age and Growth: Larvae roughly 4.62-5.09 mm (0.18-0.20 in) TL at hatching.6 Females experience faster growth and are often larger as adults than males. Length at age-1, 74-

244 mm (2.91-9.61 in) TL.18 Optimal growth temperature 22.1°C (71.78°F).18 Capable of reaching 711.2 mm (28 in) TL and 5.44 kg (12 lbs). Longevity in its northern range is 13 years.2,17

Food and Feeding: Top-level predators. Larvae 17-80 mm (0.67-3.15 in) TL primarily feed on copepods, and larger larvae and juveniles consume copepods, cladocerans including Daphnia, and benthic invertebrates.4 Piscivory increases with increased Sauger length, and fish become a primary food source at 70-110 mm (2.76-4.33 in) TL.5 Adult diets similar to walleye, and diet overlap is consistently high in sympatric populations.8

Adults primarily piscivorous feeding on a variety of fish species such as Gizzard Shad,

Yellow Perch, Freshwater Drum, and Rainbow Smelt, but continue to consume invertebrates such as chironomids.1,2,3,8,9 Seasonal shifts in diets occur as prey abundance and availability changes.3 Invertebrates are included in the diet of all sizes of Sauger, but become less important with increasing size. 688

Literature Cited:

1. Graeb, B.D.S. 2006. Sauger population ecology in three Missouri River mainstem

reservoirs. Thesis dissertation, South Dakota State University, Brookings.

2. Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of Canada. Fish. Res.

Board of Canada, Bull. 184. 966p.

3. Mero, S.W. 1992. Food habits of walleye and sauger in Lake Sakakawea, North

Dakota. M.S. thesis, South Dakota State University, Brookings.

4. Nelson, W.R. 1968b. Embryo and larval characteristics of sauger, walleye, and

their reciprocal hybrids. Transactions of the American Fisheries Society 97:167-

174.

5. Nelson, W.R. 1968a. Reproduction and early life history of sauger, Stizostedion

canadense, in Lewis and Clark Lake. Transactions of the American Fisheries

Society 97:159-166.

6. Nelson, W.R., N.R. Hines, and L.G. Beckman. 1965. Artificial propagation of

saugers and hybridization with walleyes. The Progressive Fish-Culturist 27:216-

218.

7. Nelson, W.R. 1969. Biological characteristics of the sauger population in Lewis

and Clark Lake. No. 21. US Fish and Wildlife Service.

8. Bellgraph, B.J., C.S. Guy, W.M. Gardner, and S.A. Leathe. 2008. Competition

potential between saugers and walleye in nonnative sympatry. Transactions of the

American Fisheries Society 137:790-800.

9. Swenson, W.A. 1977. Food consumption of walleye (Stizostedion vitreum) and

sauger (S. canadense) in relation to food availability and physical conditions in 689

Lake of the Woods, Minnesota, Shagwa Lake, and western Lake Superior. Journal

of the Fisheries Research Board of Canada 34:1643-1654.

10. Billington, N., C.C. Wilson, and B.L. Sloss. 2011. Distribution and population

genetics of walleye and sauger. Pages 105-132 in B.A. Barton, editor. Biology,

management, and culture of walleye and sauger. American Fisheries Society,

Bethesda, Maryland.

11. Pegg, M.A., J.B. Layzer, and P.W. Bettoli. 1996. Angler exploitation and

movements of anchor-tagged sauger in the lower Tennessee River. North

American Journal of Fisheries Management 16:218-223.

12. Nelson, W.R., and C.H. Walburg. 1977. Population dynamics of yellow perch

(Perca flavescens), sauger (Stizostedion canadense), and walleye (S. vitreum) in

four main-stem Missouri River reservoirs. Journal of the Fisheries Research

Board of Canada 34:1748-1763.

13. Bozek, M.A., T.J. Haxton, and J.K. Raabe. 2011. Walleye and sauger habitat.

Pages 133-179 in B.A. Barton, editor. Biology, management, and culture of

walleye and sauger. American Fisheries Society, Bethesda, Maryland.

14. Gangl, R.S., D.L. Pereira, and R.J. Walsh. 2000. Seasonal movements, habitat

use, and spawning areas of walleye Stizostedion vitreum and sauger S. canadense

in Pool 2 of the upper Mississippi River. Minnesota Department of Natural

Resources, Investigational Report 482, St. Paul.

15. Walburg, C.H. 1972. Some factors associated with fluctuation in year-class

strength of sauger, Lewis and Clark Lake, South Dakota. Transactions of the

American Fisheries Society 101:311-316. 690

16. Graeb, B.D.S., M.A. Kaemingk, and D.W. Willis. 2007. Early life history of

sauger in Missouri River reservoirs. South Dakota Department of Game, Fish and

Parks, Completion Report 07-08, Pierre.

17. Etnier, D.A., and W.C. Starnes. 1993. The fishes of Tennessee. University of

Tennessee Press, Knoxville.

18. Bozek, M.A., D.A. Baccante, and N.P. Lester. 2011. Walleye and sauger life

history. Pages 233-286 in B.A. Barton, editor. Biology, management, and culture

of walleye and sauger. American Fisheries Society, Bethesda, Maryland.

691

Zander, Sander vitreus (Linnaeus, 1758)

Etymology and Synonyms: Sander = “pike-perch”, referring to the genus’s resemblance to the non-related pike family (Esocidae); lucioperca = “lucio” deriving from “lucius”,

Latin for the name “pike”, and “perca”, Greek for “perch”; likely suggesting the species resemblance to both the pike family (Esocidae) and perch family (Percidae). Also commonly referred to as the Pikeperch.

Description: Body fusiform, laterally compressed and elongate. Dorsally dark olive green to gray; laterally silvery green to gray with 8-12 dark transverse bars running vertically along the body; ventrally white; dorsal and caudal fin membranes with rows of dark spots present; fins translucent olive to gray. Head moderate, conical with strongly serrated preopercle. Snout elongate with blunt tip. Eye large; placed laterally on upper portion of head; tapetum lucidum (reflective layer behind retina) present. Mouth large, terminal. Teeth large; pronounced canine teeth arranged in narrow rows on both jaws.

Dorsal fin divided into two by distinct notch; anterior dorsal fin with 13-20 spines; posterior dorsal fin with 1-2 spines and 18-24 soft rays. Adipose fin absent. Caudal peduncle thick, short. Caudal fin deeply forked with 17 soft rays and rounded distal ends on tips. Anal fin with 2-3 spines and 10-14 soft rays; insertion directly under or slightly posterior to insertion of posterior dorsal fin. Pelvic fin insertions anterior to insertion of anterior dorsal fin. Lateral line complete with 80-97 cycloid scales in series.

Similar Species: Closely resembles the Walleye, Sander vitreus, and the Sauger, Sander canadensis in the Dakotas. Walleye lack rows of dark spots on the membranes of the dorsal fin, and also display a white patch of color on the lower lobe of the caudal fin.

Sauger also have rows of dark spots present on the membranes of the dorsal fin, however 692 they appear to be more of a sandy, dull brown color with three to four, dark, oblong saddle band marks.

Distribution and Habitat: Native to coastal brackish waters, large rivers and lakes of continental Europe to western Siberia, and has the most widespread distribution of the three species of Eurasian Sander species (Zander, S. lucioperca, Volga Pikeperch, S. volgensis, and the Sea Pikeperch, S. marinus.1,2,5 Throughout Eurasia, Zander have also been introduced to Spain, the UK, France, the Netherlands, western Germany, Denmark,

Italy, Lithuania, Latvia, and Turkey as a means of commercial and recreational fishing.2,5

In 1989, 180,000 Zander fry and 1,050 fingerlings were transported from Europe to North

America, and stocked as a one-time effort into Spiritwood Lake in Stutsman Country,

North Dakota in hopes of establishing a valued recreational fishery.2 Spiritwood Lake was chosen as the experimental site of introduction due to the waterbody being completely enclosed. Following the initial stocking, the effort to continue working towards establishing a Zander fishery in Spiritwood Lake for the years to come was terminated due to concerns of the species being invasive on native fish populations. Since the stocking of Zander in Spiritwood Lake, catches of the species have been rare; however there is a small natural reproducing population that is known from angler reports and the occasional captures of age-2 adults and young-of-the-year from sampling events conducted by the North Dakota Game and Fish Department.1 Zander are known to thrive in moderately eutrophic, turbid waters with high levels of dissolved oxygen.5

Reproduction: Spawning takes place at dawn or night in the spring (generally April through May) when water temperatures reach 10-14 °C (50.0-57.2 °F) in lakes and rivers with sand, gravel, or clay substrates.3,5 Spawning migrations into spawning sites begin 693 one month prior to spawning, and consist of movements generally 10-30 km (6.21-18.64 mi), but may be up to 250 km (155.34 mi).4,5 In Europe, individuals foraging in brackish waters move inland to freshwater to spawn.5 Males construct shallow nests about 0.5 m

(19.7 in) in diameter and 5-10 cm (2.0-3.9 in) deep within sand or stone substrates, and provide parental care by guarding and fanning the eggs and fry.4 Spawning is monogamous, with a single female releasing all of her eggs at once with a single male.2

Fecundity varies greatly with the size of the female, and is known to be influenced by food supply.4 Eggs adhesive to exposed plant roots within the nest, and average 0.9 mm

(0.04 in) in diameter.4 Sexual maturity reached at age 2-5, with age at sexual maturity varying between populations due to varying growth rates.4,5 Optimal egg incubation requires water temperatures of 12-20 °C (53.6-68.0 °F).4,5 Hatching occurs at roughly 110 degree days.5

Age and Growth: Larvae roughly 4-5 mm (0.16-0.20 in) at hatching.5 Length-at-age data scarce for the small population in Spiritwood Lake, although one large individual from the population measured roughly 889 mm (35 in), and weighed 7.22 kg (15.91 lbs.).

Capable of reaching 1300 mm (51.18 in) and 20 kg (44 lbs.).5 In Europe, individuals from northern populations generally experience slower growth and reach 20-24 years of age, while individuals from the southern parts of the fishes distribution experience increased growth rates and only reach 8-9 years of age.4,5

Food and Feeding: Larvae forage on small zooplankton. Fry between the lengths of 10 and 25 mm (0.39-0.98 in) start to exhibit small amounts of piscivore behavior. Once individuals reach 100 mm (3.94 in), the diet is almost entirely fish.5 Adults are top predator pelagic piscivores, and feed heavily on prey of smaller sizes such as pelagic 694 cyprinids and smelt species. Due to this, Zander have been considered pest species in some areas, and have also been used as a biomanipulation tool in order to reduce the number of unwanted fish in some waters.4

Literature Cited:

1. Fuller, P. 2011. Sander lucioperca. USGS Nonindigenous Aquatic Species

Database, Gainesville, FL.

http://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=830 Revision Date:

10/11/2017.

2. Haponski, A.E., and C.A. Stepien. 2013. Phylogenetic and biogeographical

relationships of the Sander pikeperches (Percidae: Perciformes): patterns across

North America and Eurasia. Biological Journal of the Linnean Society, 110:156-

179.

3. Kottelat, M., and J. Freyhof. 2007. Handbook of European freshwater fishes.

Publications Kottelat, Cornol, Switzerland. 646 p.

4. Lappalainen, J., H Dörner, and K. Wysujack. 2003. Reproduction biology of

pikeperch (Sander lucioperca (L)) – a review. Ecology of Freshwater Fish 12:95-

106.

5. Larsen, L.K., and S. Berg. 2014. NOBANIS – Invasive alien species fact sheet-

Stizostedion lucioperca-from: online database of the European Netwrok on

Invasive Alien Species-NOBANIS. http://www.nobanis.org. Date of access

10/30/2019.

695

Walleye, Sander vitreus (Mitchill, 1818)

Etymology and Synonyms: Sander = “pike-perch”, referring to the genus’s resemblance to the non-related pike family (Esocidae); vitreum = “glassy”, referring to the reflective, glassy looking eyes.

Description: Body fusiform, elongate, cylindrical in cross section but slightly laterally compressed. Dorsally brown, golden brown, to olive; laterally golden brown to olive with small, speckled dark spots; ventrally cream to white; anterior dorsal fin with no distinct horizontal spots, and posterior membranes with dark pigment; posterior dorsal fin with rows of small, dark spots; lower lobe of caudal fin with white tip. Head large, elongate.

Snout conical, blunt. Cheek with little to no scales. Eye large, placed laterally on head; tapetum lucidum (reflective layer behind retina) present. Mouth large, terminal, oblique; upper jaw extends past pupil of eye; jaws nearly equal in length, with lower jaw occasionally slightly extending past upper jaw. Teeth large, sharp, canine, present on both jaws. Gill rakers long, thin, roughly 8-11. Dorsal fin with two distinctly separate fins; anterior fin with 13-14 spines; posterior fin with 1-2 spines and 19-22 soft rays. Adipose fin absent. Caudal peduncle elongate, thick. Caudal fin forked. Anal fin with 2 spines and

12-14 rays; insertion posterior to insertion of posterior dorsal fin. Pelvic fins thoracic.

Pectoral fins with 13-16 rays. Lateral line complete with 77-90 ctenoid scales in series.

Spawning males similar to non-spawning adults. Juveniles more elongate with dark, dusky bands on dorsal and lateral sides.

Similar Species: Closely resemble and occur sympatrically with Sauger. Sauger have a distinctly scaled check, a spotted anterior dorsal fin lacking dark pigment on posterior membranes, and no white tip on lower lobe of caudal fin. 696

Distribution and Habitat: Native to much of Canada, and throughout most of the northern United States from the Dakotas in the west, east throughout the Missouri,

Mississippi, St. Lawrence-Great Lakes, and Arctic River drainages to New York, and south to Arkansas and Alabama. Due to the species popularity as a sportfish, its distribution is widely expanded due to introductions and stockings. A cool-water species, with an optimal water temperature of 20-24°C (68-75.2°F).2 Walleye are pelagic until 25-

30 mm (0.98-1.18 in) TL, when they begin to reside at deeper depths near the bottom.4

Capable of tolerating a wide variety of environmental conditions.2 Preferred habitat in late spring, summer, and early autumn includes cool, clear, deep, and dark open-waters of large lakes, reservoirs, and rivers over sand, gravel, cobble, or rocky substrates.2,10

Individuals moving into dark, deep pools during the warmest days in summer (>30°C

(86°F)) often do so to seek cooler temperatures or higher dissolved oxygen concentrations.10 Mortality known to occur when water temperatures rise above 32°C

(89.6°F).7 Activity decreases around 5°C (41°F), when fish occupy overwintering habitat consisting of deep pools or sheltered areas with little current and turbulance.10 Prefer dissolved oxygen levels above 5.0 mg/L.2

Reproduction: Spawning migrations take place days before spawning begins, and are often extensive.10 Spawning takes place at night soon after ice out, or when water temperature reaches 7-10°C (44.6-50°F).3 Spawning grounds consist of shallow, wave washed shoreline areas in lakes, and flowing water in rivers with rocky substrate.2 No nests are constructed, and no parental care is given. Broadcast spawners. Fecundity

20,000-80,000 eggs/kg (9,000-36,000 eggs/lb) of body weight.17 Eggs roughly 2 mm

(0.08 in) in diameter and adhesive.3,4 Hatching takes place within 7 days in water 13.9°C 697

(57.02°F), but may take up to 14 days depending on water temperature.4,24 Hatching success varies depending on spawning date, but increases with maternal age.18

Reproductive success of the Walleye within its native range is highly variable, as exemplified in the Dakotas where populations are managed with regular fry and fingerling stockings.19,20,21 Annual recruitment dependent on factors such as fluctuating water levels during spring, discharge, wind, and maximum winter temperature.3,7,11,13

Water temperatures below 10°C (50°F) are needed for adult walleye gonad maturation, and if winter temperatures are not cold enough for a duration of time, spawning may be skipped that year causing low recruitment.12

Age and Growth: Larvae roughly 6.0-8.6 mm (0.24-0.34 in) TL at hatching. Majority of annual growth occurs from mid-June to August in northern populations.6,7 Mean relative weight known to increase during high-water periods.14 Females often larger than males.

Statewide mean lengths-at-age from South Dakota are reported as: age-1, 169 mm (6.65 in) TL; age-2, 279 mm (10.98 in) TL; age-3, 360 mm (14.17 in) TL; age-4, 425 mm

(16.73 in) TL; age-5, 490 mm (19.29 in) TL.1 Capable of reaching 1066.8 mm (42 in) TL and 8.16 kg (18 lbs). Average longevity 5-7 years; capable of reaching 18 years.4

Food and Feeding: Opportunistic feeders. Feeding mainly takes places at night. Feeding habits influenced by fluctuating water levels and temperature.8 Age-0 walleye prefer prey fishes approximately 33% of their total length, but are capable of consuming spinous prey fishes approximately 40% of their total length.22,23 Larvae 8-9 mm (0.31-0.35 in) TL primarily feed on small zooplankton such as cladocerans and copepods.5 Pisciviory increases with increased Walleye length.15,25 Walleye as small as 20-40 mm (0.79-1.57 in) TL are known to consume fish, and in an eastern South Dakota lake, Walleyes >62 698 mm (2.44 in) TL included Fathead Minnows in their diet, and Walleyes >106 mm (4.17 in) TL primarily fed on Fathead Minnows, indicating a shift to piscivory.5,16 Adults mainly piscivorous, consuming species such as Yellow Perch, White Bass, and Fathead

Minnows as well as Gizzard Shad and Rainbow Smelt in Missouri River impoundments, but continue to prey upon crayfish, frogs, and macroinvertebrates such as Diptera larvae.8,9,15,25,26

Literature Cited:

1. Willis, D.W., D.A. Isermann, M.J. Hubers, B.A. Johnson, W.H. Miller, T.R. St.

Sauver, J.S. Sorensen, E.G. Unkenholz, and G.A. Wickstrom. 2001. Growth of

South Dakota fishes: A statewide summary with means by region and water type.

South Dakota Department of Game, Fish, and Parks, Special Report 01-05, Pierre.

2. Bozek, M.A., T.J. Haxton, and J.K. Raabe. 2011. Walleye and sauger habitat.

Pages 133-179 in B.A. Barton, editor. Biology, management, and culture of

walleye and sauger. American Fisheries Society, Bethesda, Maryland.

3. Colby, P.J., R.E. McNichol, and R.A. Ryder. 1979. Synopsis of biological data on

the walleye Stizostedion v. vitreum. FAO Fisheries Synopsis 119, Rome, Italy.

4. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison.

5. Chipps, S.R., and B.D.S. Graeb. 2011. Feeding ecology and energetics. Pages

303-319 in B.A. Barton, editor. Biology management, and culture of walleye and

sauger. American Fisheries Society, Bethesda, Maryland.

6. Kelso, J.R.M., and F.J. Ward. 1972. Vital statistics, biomass, and seasonal

production of an unexploited walleye (Stizostedion vitreum vitreum) population in 699

West Blue Lake, Manitoba. Journal of the Fisheries Research Board of Canada

29:1043-1052.

7. Carlander, K.D. 1997. Handbook of freshwater fishery biology, Vol. III. Iowa

State University Press, Ames.

8. Bryan, S.D., T.D. Hill, S.T. Lynott, and W.G. Duffy. 1995. The influence of

changing water levels and temperatures of the food habits of Walleye in Lake

Oahe, South Dakota. Journal of Freshwater Ecology 10:1-10.

9. Graeb, B.D.S., S.R. Chipps, D.W. Willis, J.P. Lott, R.P. Hanten, W. Nelson-

Stastny, and J.W. Erickson. 2008. Walleye response to rainbow smelt population

decline and liberalized angling regulations in a Missouri River reservoir. Pages

275-292 in Allen, M.S., S. Sammons, and M.J. Maceina, editors. Balancing

fisheries management and water uses for impounded river systems. American

Fisheries Society, Bethesda, Maryland.

10. Paragamian, V.L. 1989. Seasonal habitat use by walleye in a warmwater river

system, as determined by radio telemetry. North American Journal of Fisheries

Management 9:392-401.

11. DeBoer, J.A., K.L. Pope, and K.D. Koupal. 2013. Environmental factors

regulating the recruitment of walleye Sander vitreus and white bass Morone

chrysops in irrigation reservoirs. Ecology of Freshwater Fish 22:43-54.

12. Colby, P.J., and S.J. Nepszy. 1981. Variation among stocks of walleye

(Stizostedion vitreum vitreum): management implications. Canadian Journal of

Fisheries and Aquatic Sciences 38:1814-1831. 700

13. Quist, M.C., C.S. Guy, and J.L. Stephen. 2003b. Recruitment dynamics of

walleyes (Stizostedion vitreum) in Kansas reservoirs: generalities with natural

systems and effects of a centrarchid predator. Canadian Journal of Fisheries and

Aquatic Sciences 60:830-839.

14. Dembkowski, D.J., S.R. Chipps, and B.G. Blackwell. 2014. Response of walleye

and yellow perch to water-level fluctuations in glacial lakes. Fisheries

Management and Ecology 21:89-95.

15. Slipke, J.W., and W.G. Duffy. 1997. Food habits of walleye in Shadehill

Reservoir, South Dakota. Journal of Freshwater Ecology 12:11-17.

16. Walker, R.E., and R.L. Applegate. 1976. Growth, food, and possible ecological

effects of young-of-the-year walleyes in a South Dakota prairie pothole. The

Progressive Fish Culturist 38:217-220.

17. Bozek, M.A., D.A. Baccante, and N.P. Lester. 2011. Walleye and sauger life

history. Pages 233-301 in B.A. Barton, editor. Biology, management, and culture

of walleye and sauger. American Fisheries Society, Bethesda, Maryland.

18. Johnston, T.A., M.D. Wiegand, W.C. Leggett, R.J. Pronyk, S.D. Dyal, K.E.

Watchorn, S. Kollar, and J.M. Casselman. 2007. Hatching success of walleye

embryos in relation to maternal and ova characteristics. Ecology of Freshwater

Fish 16:295-306.

19. Ellison, D.G., and W.G. Franzin. 1992. Overview of the symposium on walleye

stocks and stocking. North American Journal of Fisheries Management 12:271-

275. 701

20. Grote, J.D., M.R. Wuellner, D.J. Dembkowski, B. G. Blackwell, and D.O.

Lucchesi. 2015. Evaluating factors that affect recruitment of young-of-year

walleye in eastern South Dakota natural lakes. South Dakota Department of

Game, Fish and Parks, Completion Report 16-02, Pierre.

21. Lucchesi, D.O. 1997. Evaluation of large walleye fingerling stocking in eastern

South Dakota lakes. South Dakota Department of Game, Fish and Parks,

Completion Report 97-18, Pierre.

22. Einfalt, L.M., and D.H. Wahl. 1997. Prey selection by juvenile walleye as

influenced by prey morphology and behavior. Canadian Journal of Fisheries and

Aquatic Sciences 54:2618-2626.

23. Knight, R.L., F.J. Margraf, and R.F. Carline. 1984. Piscivory by walleyes and

yellow perch in western Lake Erie. Transactions of the American Fisheries

Society 113:677-693.

24. Cross, F.B. 1967. Handbook of fishes in Kansas. University of Kansas Museum

of Natural History, Miscellaneous Publication Number 45:1-357.

25. Starostka, A.B. 1999. Food habits and diet overlap of age-1 and older walleye and

white bass in Lake Poinsett, South Dakota. M.S. thesis, South Dakota State

University, Brookings.

26. Wuellner, M.R. 2009. Exploring competitive interactions between walleye and

smallmouth bass in South Dakota waters. Thesis dissertation, South Dakota State

University, Brookings.

702

CHAPTER 23

FAMILY SCIAENIDAE

Introduction

The drum and croaker family, Sciaenidae, consists of over 280 species worldwide, with roughly 80 species occurring in North America. The large family is comprised of almost entirely marine species, with many occurring throughout the tropical and coastal waters of the Atlantic, Indian, and Pacific oceans. However, a few number of freshwater species occur in South America, but only one, the Freshwater Drum (Aplodinotus grunniens) inhabits North America. Considered a warm-water species of fish, Freshwater

Drum originated from the Gulf of Mexico, before they became tolerant of entirely freshwater systems. Over time, they moved further inland up the Mississippi River and its major tributaries, eventually reaching North Dakota and South Dakota.

Drums and croakers are characterized by having a complete lateral line that reaches to the end of the caudal fin, one or two anal fin spines, and a dorsal fin that is usually separated into two parts by a deep notch. Internally, the family is well known for their rather large ear bones, called otoliths. The families common names, ‘drums’ and

‘croakers’ comes from the often loud, drumming, grunting and croaking noises that are emitted during the spawning season, or when they are taken out of the water. Contracting muscles vibrating against the swim bladder create the noises. Drums and croakers also have unusually large and robust pharyngeal jaws and teeth that help crush mollusks, which play an important part in adult’s diets, along with crayfish and fish. 703

Freshwater Drum in the Dakotas mainly occur in large turbid rivers in areas with low current, however they are also found within streams, lakes and impoundments over mud and silty substrates. They are pelagic, broadcast spawners and have extremely high fecundity rates. Although Freshwater Drum are considered a rough fish in the Dakotas and are not a desired species, certain marine members of the family can reach large sizes, making them great sportfish and important commercial fish.

704

Freshwater Drum, Aplodinotus grunniens (Rafinesque, 1819)

Etymology and Synonyms: Aplodinotus = Greek for “single back”, possibly referring to the elongate dorsal fin; grunniens = “grunting”, referring to the sound made when muscles contract against the swim bladder. Other common names include Sheephead,

Grunter, and Gray Bass.

Description: Body deep, strongly dorsolaterally compressed with concave, arched, steep profile anterior to dorsal fin insertion. Dorsally bronze to olive brown; laterally light brown to silver with iridescent sheen; ventrally silver to white; pectoral and pelvic fins light gray to white with remaining fins dusky gray-brown. Head small, conical, fully scaled. Opercle scaled. Eye large, placed laterally on top half of head. Snout rounded, slightly extending past subterminal mouth. Upper jaw extends to or past middle of eye.

Barbels absent. Lips fleshy, smooth. Teeth conical, small, present on upper and lower jaws; pharyngeal arches large, and contain fused cardiform, villiform, and molariform teeth forming a solid surface. Dorsal fin elongate with two lobes connected by shallow membrane; 8-10 spines anteriorly followed by 1 spine and 24-32 rays. Adipose fin absent. Caudal peduncle thin, short. Caudal fin rounded. Anal fin with 2 spines, second spine elongate followed by 7 rays. Pelvic fins thoratic with 1 spine and 5 rays; first ray elongate into filament. Pectoral fin with 17-18 rays. Lateral line complete, arched upward extending through caudal fin with 48-53 ctenoid scales in series. Spawning adults similar to non-breeders. Juveniles similar to adults with more triangular shaped caudal fin.

Similar Species: Does not closely resemble any other species of fish in the Dakotas. May be mistaken for carpsuckers (Carpiodes sp.) or buffalos (Ictiobus sp.) because of the deep 705 body; however, these species lack spines in the dorsal and anal fins, and lack a rounded caudal fin.

Distribution and Habitat: Only freshwater member of the family Sciaenidae.

Transitioned from brackish water in the Gulf of Mexico before becoming tolerant of freshwater, moving inland up the Mississippi River and its major tributaries.1 Native to

North America east of the Rocky Mountains from Montana and southern Saskatchewan in the west, east throughout the Missouri, Mississippi, Great Lakes, and Hudson Bay drainages to Quebec and New York, and south to the Gulf of Mexico. Largest latitudinal range of any North American freshwater fish.2,7 Abundant throughout the Upper

Mississippi River and its major tributaries. Primarily a large river species inhabiting areas with low current, but also occurs in streams, lakes, and impoundments. Prefers areas with warm, turbid water and mud substrate. Larvae pelagic, and migrate from the surface to deeper water after reaching 10 mm (0.39 in) TL, and migrate to floodplain habitat once a size of 20-35 mm (0.79 in-1.38 in) TL is attained.11

Reproduction: Pelagic, broadcast spawners. Spawning takes place in the spring when water temperatures reach 18-24.5°C (64.4-76.1°F).11 Sexual maturity occurs at age 3-4.7

Males likely attract females by producing grunting, or dumming sounds by contracting muscles against the swim bladder during spawning period, similar to when they are taken out of the water.9 Spawning period lasts 6-7 weeks, with females likely spawning more than once.11 Extremely high fecundity with females capable of producing 34,000-1 million eggs or greater.11,7 Eggs semi-buoyant often seen floating near the surface, transparent with single oil droplet, and roughly 1.48 mm (0.06 in) in diameter.11 Hatching occurs 1-2 days after fertilization. 706

Age and Growth: Larvae roughly 3 mm (0.12 in) TL at hatching.14 Growth of juveniles rapid. Mean lengths-at-age from Lewis and Clark Lake, South Dakota are reported as: age-1, 91 mm (3.58 in) TL; age-2, 166 mm (6.54 in) TL; age-3, 216 mm (8.50 in) TL; age-4, 252 mm (9.92 in) TL; age-5, 285 mm (11.22 in) TL; age-6, 308 mm (12.13 in) TL; age-7, 325 mm (12.80 in) TL.12 Commonly 304.8-508 mm (12-20 in) TL, but were once capable of reaching 711 mm (4 ft) TL and kg (60 lbs).15 Long-lived species. Average lifespan 8-10 years; longevity 72 years.6 Females generally larger and have higher growth rates than males.7

Food and Feeding: Opportunistic omnivore. Feeding takes place day and night. Diet shift from soft to hard prey occurs with increasing size.4,5,13 Larvae highly dependent on zooplankton such as copepods and cladocerans, and grow to select larger prey as individual size and gape limitations increase.8,10,11 Larval Freshwater Drum may have the potential to compete for desired zooplankton with larval sportfish species.10 Juveniles feed on chironomids, mayfly and caddisfly larvae. Adults consume a wide variety of prey and primarily suction feed along the bottom on fish, mollusks, and crayfish.3,4 Exhibit diurnal food habits, consuming zooplankton during the day and benthic organisms during the night.12 Primary prey items known to change throughout the year, with adults consuming a greater number of fish August-November.5 Robust pharyngeal jaws and molariform teeth assist adults with crushing mollusks.

Literature Cited:

1. Barney, R.L. 1926. The distribution of the freshwater sheephead, Aplodinotus

grunniens, in respect to the glacial history of North America. Ecology 7:351-364. 707

2. Boschung, H.T., Jr., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian

Books, Washington, D.C.

3. Diaber, F.C. 1950. The life history and ecology of the sheephead, Aplodinotus

grunniens Rafinesque, in western Lake Erie. Ph.D. dissertation, Ohio State

University, 150pp.

4. Essner, R.L. Jr., R. Patel, and S.M. Reilly. 2014. Ontogeny of body shape and diet

of freshwater drum (Aplodinotus grunniens). Transactions of the Illinois State

Academy of Science 107:27-30.

5. Griswold, B.L., and R.A. Tubb. 1977. Food of Yellow Perch, White Bass,

Freshwater Drum, and Channel Catfish in Sandusky Bay, Lake Erie. Ohio Journal

of Science 77:43-47.

6. Pereira, D.L., Y. Cohen, and G.R. Spangler. 1992. Dynamics and species

interactions in the commercial fishery of the Red Lakes, Minnesota. Canadian

Journal of Fisheries and Aquatic Sciences 49:293-302.

7. Rypel, A.L. 2007. Sexual dimorphism in growth of freshwater drum. Southeastern

Naturalist 6:333-342.

8. Schael, D.M., L.G. Rudstam, and J.R. Post. 1991. Gape limitation and prey

selection in larval yellow perch (Perca flavescens), freshwater drum (Aplodinotus

grunniens), and black crappie (Pomoxis nigromaculatus). Canadian Journal of

Fisheries and Aquatic Sciences 48:1919-1925.

9. Schneider, H., and A.D. Hasler. 1960. Laute und Lauterzeugung beim

Süsswassertrommler Aplodinotus grunniens Rafinesque (Sciaenidae, Pisces).

Zeitschrift für vergleichende Physiologie 43:499-517. 708

10. Sullivan, C.L., K.D. Koupal, W.W. Hoback, B.C. Peterson, and C.W.

Schoenebeck. 2012. Food habits and abundance of larval freshwater drum in a

South Central Nebraska irrigation reservoir. Journal of Freshwater Ecology

27:111-121.

11. Swedberg, D., and C. Walburg. 1970. Spawning and early life history of the

freshwater drum in Lewis and Clark Lake, Missouri River. Transactions of the

American Fisheries Society 99:560-570.

12. Swedberg, D.V. 1968. Food and growth of the freshwater drum in Lewis and

Clark Lake, South Dakota. Transactions of the American Fisheries Society

97:442-447.

13. Wahl, D.H., K.A. Bruner, and L.A. Nielsen. 1988. Trophic ecology of Freshwater

Drum in large rivers. Journal of Freshwater Ecology 4:483-491.

14. Walburg, C.H. 1976. Changes in the fish populations of Lewis and Clark Lake,

Missouri River, 1956-1962. U.S. Fish and Wildlife Service Special Report,

Fisheries No.482. 27pp.

15. Witt, A. Jr. 1960. Length and weight of ancient freshwater drum, Aplodinotus

grunniens, calculated from otoliths found in Indian Middens. Copeia 3:181-185.

709

CHAPTER 24

FAMILY CICHLIDAE

Cichlids, are the common name for the group of fishes from the extremely diverse and one of the largest vertebrate families, Cichlidae. Species of the family are native to the warm, tropical, and temperate waters of Africa, Asia, Central and South America.

Due to belonging to such a large family, the variations in body size amongst the numerous species is very diverse, however body morphology is relatively similar and can primarily be split into two main groups: deeply laterally compressed or cylindrical and elongate. Cichlids can be recognized by having a single nostril on each anterior lateral side of the head, and a lateral line divided into an anterior upper section ending below the posterior end of the dorsal fin base, and a posterior lower section extending to the center of the caudal peduncle. All species within the family Cichlidae also share the trait of having fused lower pharyngeal bones, however there is considerable variation amongst the tooth patterns and shapes. Cichlids also have highly movable jaws, and this trait combined with the variations in tooth patterns and shapes allow Cichlids to efficiently feed, capture and process a wide variety of prey items. The majority of Cichlids feed on plant matter and a variety of small invertebrates.

Cichlids often times show aggressive behavior during the spawning period, as males are known to establish territories and develop vivid colors to assist in the domination over other males. Most Cichlids spawn monogamously or polygamously, and exhibit parental care, which primarily consists of protecting the eggs and fry from predators. Some species of Cichlids even practice mouthbrooding, which entails either one or both parents guarding the eggs or larvae orally. The bright coloration and unique 710 behaviors make Cichlids desired and popular aquarium species. Larger species of

Cichlids are also desired and highly sought after food and game fish. The group of

Cichlids called Tilapiines, better known as Tilapia, are a fast growing, low cost species that is important to the aquaculture industry and are one of the most popular consumed groups of fishes in many parts of the world, including the United States. An example of highly sought after sport fish within the family Cichlidae are the Peacock bass from the genus Cichia, which are native to South America but have been introduced to the United

States in southern Florida.

Only one species of Cichlid occurs in the Dakotas. The nonnative Jack

Dempsey, Rocio octofasciata, has established populations in the Fall River as well as in the warmer waters of southern Black Hills streams in western South Dakota, and was likely introduced by purposeful or accidental aquarium release.

711

Jack Dempsey, Rocio octofasciata (Regan, 1903)

Etymology and Synonyms: Rocio = named after the author Charles T. Regan’s wife,

Rocio, a Spanish name meaning “morning dew”, in reference to the species numerous electric blue-cyan spots; octofasciata = Latin for “octo-”, meaning “eight”, and “-fascia”, meaning “a belt or a stripe”, referring to the 8-11 dark, diffuse vertical bars or stripes on lateral sides of young and juvenile individuals. The species common name refers to the

1920’s famous boxer, Jack Dempsey, in relation to the species aggressive behavior.

Description: Body elongate, moderately deep. Dorsally gray-olive, dark blue-green to black; laterally olive to brown with 8-11 dark, diffuse vertical bars or stripes (most prominent in preserved specimens); ventrally dark brown to olive; head, body, and unpaired fins with numerous electric blue-cyan spots; bright to deep red coloration, especially on the caudal and edge of the dorsal fin; two noticeable dark spots, one on the middle of the body, with the second on the posterior end of the caudal peduncle; adult males develop dark, horizontal stripes on the forehead. Head large. Snout conical, short.

Eye moderately large, placed laterally on head; iris gray to bronze. Mouth terminal; upper jaw barely extends to anterior edge of eye. Lips fleshy. Dorsal fin elongate with 17-19 spines followed by 9-10 rays. Caudal peduncle short, thick. Caudal fin with slightly rounded distal end. Anal fin elongate with 8-9 spines followed by 7-8 rays forming a short, pointed filament on posterior end. Pelvic fins thoracic. Pectoral fins with rounded distal end. Spawning males generally darker in color, almost black in appearance.

Juveniles with more prominent vertical bars or stripes on lateral sides, and less numerous electric blue-cyan spots. 712

Similar Species: No other species within the Dakotas exhibit similar features to the Jack

Dempsey.

Distribution and Habitat: Only species from the large and diverse family Cichlidae that occurs within the Dakotas. Cihchlids are native to warm, tropical, and temperate waters of Africa, Asia, Central and South America, and are one of the most popular aquarium species of cichlid fish.1,9 In Central America, Jack Dempsey are native to southern

Mexico and Honduras.8 Cichlids are extremely adaptable to new ecological conditions, with the Jack Dempsey being no exception. The species has managed to effectively inhabit and become established in areas considerably north of its native range, including a population in Fall River, South Dakota, adjacent to the town of Hot Springs, as well as in warmer waters of southern Black Hills streams.9,12 Means of introduction likely due to purposeful or accidental aquarium release. In the wild, the species prefers to inhabit areas with warm, clear, to slightly murky waters such as small spring fed lakes and shallow rivers with little to no current, and streams with large amounts of aquatic vegetation over soft, muddy or sand substrate.8 Inhabits water bodies within its native range where water temperature is 22-30°C (71.6-86°F).5 Loss of equilibrium occurs at 9°C (48.2°F), and lower lethal water temperature is reported as 8°C (46.4°F).11 Able to withstand low oxygen concentrations.7 Not capable of occupying waters where salinity exceeds 8%.2

Reproduction: Little information available on spawning in the wild. Sexual maturity reached at 75 mm (2.95 in) TL.1,6 Fecundity does not usually exceed 800 eggs per female, but has been reported as 1010 eggs per female.9,10 Like many other species in the family

Cichlidae, parental care and protection from predators is given to eggs and fry by one or both parents.4 Both parents known to increase the availability of food and feeding 713 opportunities for their fry by “fin-digging”, or stirring up the bottom substrate with their pectoral fins in order to release detritus, zoo- and phyto- benthos.13,14 Exhibits aggressive and unpretentious behavior.14

Age and Growth: Juveniles vary 38-59 mm (1.50-2.32 in) SL.9 Average length 127-203 mm (5-8 in) TL. Capable of reaching roughly 200-250 mm (7.87-9.84 in) TL.1,3,6

Longevity 10 years.

Food and Feeding: Omnivore. Adults actively feed on a variety of plant and animal matter. Main prey items include chironomids, trichopterans, and odonata larvae, as well as aerial insects, small fish, gastropods, and eggs of other aquatic invertebrates.9

Sunflower seeds have also been identified in adult diets, indicating flexibility and adaptation to consume prey that is not generally consumed, but abundant and accessable.9

Foraging activity decreases at 16°C (60.8°F), and ceases at 13°C (55.4°F).11 Little information available on juvenile diets.

Literature Cited:

1. Cadwallader, P.L., G.N. Backhouse, and R. Fallu. 1980. Occurrence of exotic

tropical fish in the cooling pondage of a power station in temperate southeastern

Australia. Australian Journal of Marine Freshwater Resources 31:541-546.

2. Dial, R.S., and S.C. Wainright. 1983. New distributional records for non-native

fishes in Florida. Florida Scientist 46:8-15.

3. Fuller, P.L., L.G. Nico, and J.D. Williams. 1999. Nonindigenous fishes

introduced into the inland waters of the United States. Special Publication 27.

American Fisheries Society, Bethesda, Maryland. 714

4. Keenleyside, M.H.A. 1991. Cichlid fishes: behavior, ecology and evolution.

Chapman and Hall, London, New York.

5. Konings, A. 1989. Cichlids from Central America. T.F.H. Publications, Neptune

City, New Jersey.

6. Legge, R. 1970. The complete aquarists guide to freshwater tropical fishes.

Eurobook Limited, London, England.

7. Obordo, C.O., and L.J. Chapman. 1997. Respiratory strategies of a non-native

Florida cichlid, Cichlasoma octofasciatum. Florida Scientist 60:40-52.

8. Page, L.M., and B.M. Burr. 1991. A field guide to freshwater fishes of North

America north of Mexico. Houghton Mifflin Company, Boston.

9. Pashkov, A.N., and D.D. Zvorykin. 2009. Some morphological specific features

of cichlasomine Rocio octofasciata (Perciformes, Cichlidae) from the population

in Lake Staraya Kuban. Journal of Ichthyology 49:383-389.

10. Sakurai, A., Y. Sakamoto, and F. Mori. 1993. Aquarium fish of the world: the

comprehensive guide to 650 species. Chronicle Books, San Francisco.

11. Shafland, P.L., and J.M. Pestrak. 1982. Lower lethal temperatures for fourteen

non-native fishes in Florida. Environmental Biology of Fish 7:149-156.

12. Simpson, G., J. Carreiro, G. Galinat, J. Davis, B. Miller, C. Pasbrig, B. Fletcher,

M. Barnes, D. Jones, and M. Bucholz. 2014. Black Hills fisheries management

area strategic plan 2014-2018. South Dakota Game, Fish and Parks Wildlife

Division. 715

13. Zworykin, D.D. 1998. Parental fin digging by Cichlasoma octofasciatum

(Teleostei: Cichlidae), and the effect of parents’ satiation state on brood

provisioning. Ethology 104:771-779.

14. Zworykin, D.D., and A.N. Pashkov. 2010. Eight-striped cichlasoma-an

allochthonous species of cichlid fish (teleostei: cichlidae) from Staraya Kuban

Lake. Russian Journal of Biological Invasions 1:1-6.