Spawning Habitat Characteristics Of Great Lakes Fishes J.A. Lane1, C.B. Portt1 and C.K. Minns2 1. C. Portt and Associates, 56 Waterloo Avenue, Guelph, Ontario, Canada N1H 3H5 2. Department of Fisheries and Oceans, Great Lakes Laboratory for Fisheries and Aquatic Sciences, Bayfield Institute, 867 Lakeshore Road, PO Box 5000, Burlington, Ontario, L7R 4A6 Canada October 1996 Canadian Manuscript Report of Fisheries and Aquatic Sciences No. 2368 Fisheries Pêches and Oceans et Océans Canadian Manuscript Report of Fisheries and Aquatic Sciences Manuscript reports contain scientific and technical information that contributes to existing knowledge but which deals with national or regional problems. Distribution is restricted to institutions or individuals located in particular regions of Canada. However, no restriction is placed on subject matter, and the series reflects the broad interests and policies of the Department of Fisheries and Oceans, namely, fisheries and aquatic sciences. 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Les demandes de rapports seront satisfaites par l’établissement auteur don’t le nom figure sur la couverture et la page du titre. Les rapports épuisés seront fournis contre rétribution par des agents commerciaux. CANADIAN MANUSCRIPT REPORT OF FISHERIES AND AQUATIC SCIENCES 2368 SPAWNING HABITAT CHARACTERISTICS OF GREAT LAKES FISHES by 1 1 J.A. Lane , C.B. Portt and C.K. Minns 2 1. C. Portt and Associates, 56 Waterloo Avenue, Guelph, Ontario, Canada N1H 3H5 2. Department of Fisheries and Oceans, Great Lakes Laboratory for Fisheries and Aquatic Sciences, Bayfield Institute, 867 Lakeshore Road, PO Box 5000, Burlington, Ontario, L7R 4A6 Canada © Minister of Supply and Services 1996 Cat. No. Fs97-4/2368 ISSN 0707-6473 Correct citation of this publication: Lane, J.A., C.B. Portt and C.K. Minns. 1996. Spawning habitat characteristics of Great Lakes fishes. Can. MS Rep. Fish. Aquat. Sci. 2368: v+48p. ii TABLE OF CONTENTS LIST OF TABLES .............................................................................................................................. iv ABSTRACT/RÉSUMÉ ...................................................................................................................... v 1.0 INTRODUCTION......................................................................................................................... 1 2.0 METHODS.................................................................................................................................... 2 3.0 RESULTS AND DISCUSSION................................................................................................... 3 4.0 REFERENCES............................................................................................................................. 33 APPENDIX A Great Lakes Basin Fish Species List........................................................................ 45 APPENDIX B Riverine spawners..................................................................................................... 48 iii LIST OF TABLES Table 1. Depth strata utilized and use of cover by spawning fishes. Presence within a depth stratum is indicated with an X. A dash (-) indicates that the species was not reported to utilize a particular depth stratum or cover. For species marked with an asterisk (*), inferences have been drawn, the bases for which are provided in the 'Comments' c column. The notation indicates additional information is given, also in the 'Comments' column................................................................................................................ 10 Table 2. Strength of association with substrate types, the importance of lake (versus stream) habitat . A dash (-) indicates that no information was found to indicate that the species utilizes a particular substrate type. For species marked with an asterisk (*) inferences about relationships have been drawn, the bases for which are provided in c the 'Comments' column. The notation ' ' indicates additional information is given, also in the 'Comments' column. References are listed in Table 1........................................ 17 Table 3. Season, range of water temperature and reproductive guild for Great Lakes fishes. Ranges given indicate temperatures above and below which a species will not spawn. Reproductive guilds correspond to Balon (1975) as follows: A.1 open substratum spawners = broadcast; A.2 brood hider = hide; B.2 nest spawners = nest. c Other information is indicated with the notation and details are given in the 'Comments' column. For species marked with an asterisk (*) inferences about relationships have been drawn, the bases for which are also provided in the 'Comments' column. n/a = not available................................................................................ 26 Table 4 Number of species that spawn within each depth stratum. ................................................. 32 Table 5 Number of species associated with each substrate and with cover, by strength of association.............................................................................................................................. 32 iv ABSTRACT Lane, J.A., C.B. Portt and C.K. Minns. 1996. Spawning habitat characteristics of Great Lakes fishes. Can. MS Rep. Fish. Aquat. Sci. 2368 :v+48p. A review of the literature was conducted to compile knowledge of spawning habitat characteristics of 139 species of Great Lakes fishes. Water depth, substrate, and vegetation/cover were the habitat variables examined. Nineteen Great Lakes species spawn exclusively in rivers, and one spawns in the marine environment. The majority of the remaining 120 species spawn in shallow water. Most (105) spawn in the spring. Gravel and sand are the most commonly utilized substrates. Bedrock and hard-pan clay are utilized by few species. Aquatic vegetation is an important component of spawning habitat for many species, as are other forms of cover. The majority of these species scatter their eggs, usually over a specific substrate, but 32 species guard nests, and 23 construct redds in which the eggs are concealed. The precision with which spawning habitats are selected by many species cannot be reflected with the limited number of variables which we have used to describe nearshore habitats. RÉSUMÉ J.A. Lane, C.B. Portt et C.K. Minns, 1996. Caractéristiques de la zone de frai des poissons des Grands Lacs. Can. MS Rep. Fish. Aquat. Sci. 2368 : v+48p. Une analyse documentaire a été effectuée pour rassembler les données connues sur les caractéristiques de la zone de frai des poissons des Grands Lacs. Les paramètres étudiés étaient la profondeur de l'eau, le substrat et la couverture végétale. Dix-neuf espèces pondent seulement dans les rivières, une dans un environnement marin. La majorité des 120 autres espèces déposent leurs oeufs dans l'eau peu profonde. Pour la plupart (105 espèces), le printemps est la saison du frai. Les substrats les plus fréquemment utilisés sont le gravier et le sable. Quelques espèces déposent leurs oeufs sur un substrat rocheux ou une carapace d'argile. Pour bon nombre d'espèces, la végétation aquatique, tout comme les autres formes de substrat, est une composante importante de la zone de frai. La majorité de ces espèces dispersent leurs oeufs, normalement au-dessus d'un substrat particulier. Trente-deux espèces, par contre, surveillent un nid et 23 construisent un nid de frai dans lequel elles dissimulent les oeufs. De nombreuses espèces ont des besoins particuliers en ce qui concerne la zone de frai, qui ne sont pas décrits en détail en raison du nombre limité de variables étudiées. v 1.0 INTRODUCTION The nearshore areas are crucial in providing habitat for most of the fish species in the Great Lakes, during at least one life stage. Nearshore areas are also the locations of greatest human interaction with the lakes, and include some of the most degraded habitats in the Great Lakes basin. Effective management strategies of nearshore areas must integrate the preservation of fish habitat and fish populations, and human activity which is concentrated there. 'NO NET LOSS OF THE PRODUCTIVE CAPACITY OF FISH HABITAT' is the guiding principle of the Department of Fisheries and Oceans (DFO) 'Policy for the Management of Fish Habitat' (DFO 1986). The ultimate objectives of this policy are: 1) conservation of habitat; 2) restoration of damaged habitat; and 3) development of new habitat resulting in a net gain of productive capacity (Minns et al. 1995). The difficulty in assessment of individual development projects lies in the lack of a common, quantitative protocol that can be used both by project proponents and habitat managers to assess individual projects. A prototype methodology developed by Minns et al. (1995) requires pre- and postdevelopment characterizations of fish habitats, and predictions of their relationship to fish populations and communities. These predictions are based on knowledge of spawning, nursery and adult habitat characteristics. To date, reports have been prepared documenting habitats of adult and young-of-the year fishes in the Great Lakes (Lane et al. 1996a, 1996b). This report summarizes the spawning habitat characteristics required for the defensible methods assessment process. 1 2.0 METHODS The computer database Aquatic Sciences and Fisheries Abstracts (1978-December 1995) was searched and relevant literature was examined. Secondary sources (Scott and Crossman, 1973; Becker, 1983) were relied upon for summaries of earlier literature. This report presents data for those species which spawn in the lake environment or in estuaries at the mouths of tributary streams or river deltas, which were treated as lentic habitat. As was done for nursery and adult habitats (Lane et al, 1996a,b), spawning habitat characteristics are characterized by three variables: water depth (0-1, 1-2, 2-5, 5+ metres), substrate (bedrock, boulder, cobble, rubble, gravel, sand, silt, clay, hard-pan clay), and structure/cover (emergent vegetation, submergent vegetation and other e.g. logs, rocks). The depth strata used for spawning are indicated (Table 1), but no attempt was made to assess relative importance. The estimated strength of association with various substrate materials is indicated (low, medium, high; Table 2). Substrate described as detritus or mud in the literature is included in the silt category. The significance of vegetation or other cover for spawning (Table 1) was estimated based on the available documentation. 'High' indicates a species always, or nearly always, spawns where there is a particular form of cover; 'medium' indicates a species often spawns, and 'low' indicates a species sometimes spawns where a given cover type is present. A dash (-) indicates that cover is discerned to be of no importance. When vegetation is an essential component of spawning habitat, the comments column notes that eggs adhere to vegetation. An index of the utilization of lentic, as opposed to lotic, habitat for spawning is also provided (Table 2). The range of water temperatures at which spawning occurs is provided, or if this 2 information was unavailable the temperature at which spawning peaks or commences is given (Table 3). The season during which spawning occurs is also indicated for each species (Table 3). The dates of spawning vary, due primarily to north-south climatic gradients. For the purposes of this report spring is defined as the period during which water temperatures are increasing, fall is defined as the period during which water temperatures are declining, and winter is defined as the period during which water temperatures are at their minimum, and more or less stable. The spawning behaviour of each species is broadly characterized (Table 3), based on the reproductive guilds of Balon (1975). Species are described as a) broadcast spawners which release their eggs over a given area and do not guard; b) hiders, which typically bury their eggs in redds, and do not guard; and c) nest-builders, which guard their nests. Information on spawning habitat was readily available for most fish species in the Great Lakes. Those few for which inferences were made are indicated with an asterisk (*). The basis for the inference is indicated in the 'Comments' column. 3.0 RESULTS AND DISCUSSION The species list used in this report (Appendix A) is the same as was used for the compilations of nursery and adult habitats (Lane et al, 1996a,b). This information is based on the distribution information in Mandrak and Crossman (1992) and information obtained on recently discovered exotics. No information is presented for three species which are considered extirpated from the Great Lakes (deepwater cisco, Coregonus johannae; blackfin cisco, C. nigripinnis; gravel chub, Erimystax x-punctata), or for hybrids. The American eel (Anguilla rostrata), which returns 3 to the marine environment to spawn (Tesch and Greenwood 1977), was also excluded. Information was compiled on the spawning habitat characteristics of 131 species. Inferences were drawn for an additional 8 species based on adult habitat or on the spawning habitat of species with which they are known to hybridize. Sixty-nine of the species resident in the Great Lakes spawn primarily, if not exclusively, in lentic habitats. Twenty-three species commonly utilize both lake and stream habitats for spawning; 28 species are normally stream spawners but some spawning is known to occur in the lake environment. The remaining nineteen species apparently spawn only in streams and rivers; these species are listed in Appendix B. Most fish species resident in the Great Lakes spawn on only one occasion during the year, although some species (e.g. fathead minnow, gobies, ruffe) can produce more than one brood during a breeding season. It appears that for most species, the photoperiod determines the overall reproductive state, but water temperature ultimately determines when spawning will occur. Of the species examined, 105 spawn in the spring, twelve spawn in the fall and three spawn during the winter (Table 3). Spawning dates for the same species often differ by more than a month between the upper and lower Great Lakes, and spawning dates in one location can vary by two to three weeks between years for some species, depending upon weather conditions. Sheltered shallow areas and tributaries warm more quickly in the spring than more exposed areas in the lakes. Consequently, water temperature also plays an important role in determining where some species spawn. Thirty-two of the 120 lake spawning species guard their broods (Table 3). Some of these 4 species conceal their eggs in existing cavities (i.e. brown bullhead, Ameiurus nebulosus) or under objects (i.e. fathead minnow, Pimephales promelas). Sticklebacks construct a nest of algae or plant material. In the case of the centrarchids the nest is usually only a shallow depression from which fine sediments have been fanned. All of these species were considered guarding nest spawners (guild B.2.) by Balon (1975) except white crappie (Pomoxis annularis), which he considered to be a guarding substratum chooser (guild B.1.). Based on descriptions of the spawning behaviour of white crappie, we consider its reproductive tactics to be similar to those of the other centrarchids and have placed it with them in guild B.2. Two non-guarding guilds are represented. The majority of the non-guarding species (65) broadcast their eggs rather than actively concealing them. These are, in Balon's terminology, nonguarding open-substratum spawners (guild A.1.). In most cases these eggs settle to the bottom, or onto vegetation which overlies the bottom. Exceptions include freshwater drum (Aplodinotus grunniens), whose eggs are positively buoyant and float at the surface, and yellow perch (Perca flavescens) whose eggs are contained in gelatinous strings or tubes which attach to vegetation or the bottom. Spawning habitat requirements for this group can be quite precise; for example, northern pike spawn almost exclusively over the previous year's flooded terrestrial or emergent aquatic vegetation; lake trout require shoals with sufficient wave action to remove fine sediment. For other species, such as white perch (Morone americana), neither substrate nor vegetation appear to be important. The remaining twenty-three non-guarding species either bury their eggs in redds (trouts, salmons and charrs) or clear depressions or construct mounds of pebbles over which they spawn, 5 and among which the eggs are concealed (several cyprinids). This group corresponds to Balon's (1975) non-guarding brood-hiding lithophils (guild A.2.1; the third level of the classification can be included in this case, since all utilize similar substrates). We have included two species in this guild, common shiner (Luxilus cornutus) and cutlips minnow (Exoglossum maxilingua), which Balon (1975) considered to be guarders. Although the males of both species are territorial during spawning, we interpret the descriptions of spawning behaviour to indicate that neither guards its brood. There are numerous accounts of more than one minnow species spawning over a particular area at the same time. Male common shiners defended spawning territory on most hornyhead chub (Nocomis biguttatus) redds observed by Vives (1990). Although some antagonistic encounters between the two males were observed, most of the common shiners' aggressive behaviour was directed toward other common shiners and female chubs. Rosyface shiners (Notropis rubellus) were also observed spawning over chub redds (Vives, 1990). Goff (1984) reported finding longnose gar (Lepisosteus osseus) eggs in association with a significant proportion (14%) of smallmouth bass (Micropterus dolomieu) nests. The redfin shiner (Lythrurus umbratilus) usually spawns over the nests of actively spawning sunfish especially green sunfish (Lepomis cyanellus; Hunter & Hasler 1965; Noltie 1989). Spawning activity of the shiners appears to be initiated by the odour of milt and ovarian fluid from sunfish (Hunter & Hasler 1965). In these cases, the eggs of both species are guarded by the centrarchid. More than one species defending the same spawning territory, or more than one species spawning over the same redd, could be attributed simply to selection of similar spawning habitats. An obligate symbiotic or 6 parasitic relationship apparently does not exist, as all of these species are reported to spawn independently, but such relationships may be evolving. Gravel and sand are the most commonly used substrates for spawning. Cobble and silt are also frequently utilized (Table 4), although the number of species using silt is influenced by species which spawn on vegetation, which often grows in silt substrates. Bedrock and hard-pan clay are utilized by few species (Table 4). Many species spawn on, or amongst, both emergent and submergent vegetation. Strong association with cover is also common, with emphasis on fixed objects such as rocks and logs. For a number of reasons, spawning is typically the most intensively studied aspect of a fishes' life-history. Because they move into either the nearshore areas of the lakes or the tributaries, the adults of many species are more readily observed during spawning than at any other time. They are often also most susceptible to capture during spawning or pre- and post-spawning migrations, when many individuals are concentrated in relatively small areas. The reproductive behaviour of a species is one component of an evolved 'optimum' strategy for survival, which is bounded by genetic and environmental constraints. As Kryzhanovsky (1949) stated "adaptations of fishes for spawning and development reflect not only the essential ecological factors of the embryonic period, but also the essential factors of all other intervals of life." This is true, but an individual's ability to actively select advantageous habitats and avoid those that are disadvantageous increases with age. Embryos are passive and thus at the mercy of their surroundings. Once exogenous feeding begins, a readily available food supply is essential, or death ensues. The precision of reproductive behaviour and spawning habitat selection exhibited by many 7 species reflects the vulnerability of the early life stages and the need to position them, in time and in space, where environmental conditions are most conducive to survival and growth. The characterization of spawning habitats which is provided here does not reflect factors which, for some species, are essential. For example, brook trout (Salvelinus fontinalis) select areas of upwelling for spawning (Hartley & Kelso 1991; Scott & Crossman 1973). In fact, the preference for groundwater is so pronounced that substrates of silt and detritus, or even old beaver lodges, will be utilized if seepage is present (Wetzel & MacCrimmon 1983; Webster & Eiriksdotter 1976, Fraser, 1982). Lake trout (Salvelinus namaycush) spawning shoals usually are moderately exposed to wave action and associated currents; this is attributed to the need to maintain interstices within the substrate which are free of fine sediments. In contrast, the reproductive success of smallmouth bass is negatively correlated with exposure to strong wind (Goff, 1986). The homing exhibited by many fish species may be the ultimate in precise spawning habitat 'selection'. For some species, however, precise homing could be a liability. Northern pike typically spawn in shallow water over flooded terrestrial or emergent aquatic vegetation, and the location of these habitats can vary from year to year due to water level fluctuations. There are, of course, species which appear to be true habitat generalists, such as yellow perch and white perch. As with the reports concerning adult and nursery habitat (Lane et al, 1996a,b), certain conclusions regarding the utilization of various habitats can be drawn from the information compiled, but its limitations must be noted. Our assessment of the relative significance of particular habitats to individual species was necessarily subjective. Even though spawning habitat has often received a disproportionate amount of attention from biologists, relative to other life 8 stages, descriptions of spawning habitat for some species are vague. The precision with which spawning habitats are selected by many species cannot be reflected with the limited number of variables which we have used to describe nearshore habitats. These limitations notwithstanding, the data contained in this report are useful in assessing the relative significance of various habitats to spawning fishes, particularly when applied at the guild or community level (Lane et al, 1996a; 1996b). Acknowledgements We gratefully acknowledge the constructive comments provided by J.Leslie and D. Bodaly, Department of Fisheries and Oceans; A. Mathers and S.J. Nepszy, Ontario Ministry of Natural Resources; and E. Holm, Royal Ontario Museum. 9 Table 1. Depth strata and type of cover utilized by spawning Great Lakes fishes. Presence within a depth stratum is indicated with an X. A dash (-) indicates that the species was not reported to utilize a particular depth stratum or cover. For species marked with an asterisk (*), inferences have been drawn, the bases for which are provided in the c 'Comments' column. The notation ' ' indicates additional information is given, also in the 'Comments' column. Common Name Water Depth (m) Cover References Comments 0-1 1-2 2-5 5+ Submergent Vegetation Emergent Vegetation Other lake sturgeon X X X - - - rocks, logs 11, 47, 57 adhesive eggs longnose gar X - - - high high Cladophora 5, 11, 47, 131 eggs adhere to vegetation spotted gar X - - - high high brush, debris 47, 111, 131 eggs adhere to vegetation bowfin X X - - high high - 11, 47, 136 alewife X X X - low low stones, debris 47, 75, 98, 136 eggs broadcast over substrate gizzard shad X X X - high high brush, debris 8, 47, 75, 136, 138 adhesive eggs rainbow trout X X X X - - - 1, 47, 131 Atlantic salmon X - - - - - - 1, 47, 77, 89, 131 brown trout X X - - - - - 1, 5, 47, 149 Arctic charr - X X X - - - 91, 131 brook trout X X - - - - - 24, 47, 52, 121, 144, 149 areas of upwelling lake trout X X X X - - crevices, cracks 35, 36, 47, 64, 96 depth a function of race lake whitefish X X X X - - - 1, 5, 47, 61, 131 cisco X X X X - - - 1, 47, 131 bloater - - - X - - - 1, 5, 47, 109 kiyi - - - X - - - 47, 54, 110 shortnose cisco - - - X - - - 47, 107 10 shortjaw cisco - - - X - - - 47, 58 pygmy whitefish X X - X c - - - 5, 47, 131 round whitefish X X X X - - - 1, 47, 53, 103 rainbow smelt X X X X low low - 47, 75, 128 northern pike X X - - - high flooded terrestrial vegetation 19, 47, 131, 136, 146 muskellunge X X - - high high stumps, logs 32, 33, 47, 131, 146 grass pickerel X - - - - high flooded terrestrial vegetation 22, 47 c central mudminnow X - - - high high - 47, 82, 118 c quillback X X - - - - high turbidity 1, 5, 47 longnose sucker X X X - - - - 5, 47, 131 white sucker X X - - low low - 5, 23, 47, 122, 131, 136 lake chubsucker X X - - high high filamentous algae, grass stubble 1, 47, 131 northern hogsucker X - - - - - - 5, 23, 47, 131 bigmouth buffalo X X X - high high high turbidity 43, 47, 62 spotted sucker X - - - - - depressions behind rocks 5, 23, 112 silver redhorse X - - - - - - 5, 47 shorthead redhorse X - - - - - - 5, 23, 47 greater redhorse* X X X - - - - 5, 47, 131 c c c 11 c spawning & spent caught at 31-46 m c eggs adhere to vegetation. eggs adhere to vegetation eggs adhere to vegetation *possibly spawn in lakes goldfish X X X - high - undersides of objects, roots 5, 47 northern redbelly dace X X - - high - - 1, 5, 30, 47, 131 finescale dace X X - - - - logs, brush 5, 30, 131, 135 lake chub X X - - - - under boulders 12, 47, 131 common carp X X - - high high - 47, 131, 136 cutlips minnow X - - - - - large rocks, logs 131, 139 brassy minnow* X - - - high high - 5, 131 eastern silvery minnow X - - - high high - 5, 131 silver chub X X X X - - - 5, 47, 115, 131 hornyhead chub X - - - low low - 5, 25, 83, 140 golden shiner X X - - high high organic debris, filamentous algae 14, 47, 60, 146 pugnose shiner X X - - high high - 5, 47, 76, 114 emerald shiner X X X X low low - 17, 47, 136 bridle shiner X - - - high high - 47, 51 common shiner X - - - low low - 6, 47, 60, 131, 136 often use nests of other minnows blackchin shiner* X - - - high high - 5, 47 *adult habitat blacknose shiner X - - - high high roots of aquatic vegetation 47, 131 spottail shiner X X X - medium medium - 47, 80, 136 rosyface shiner X - - - - - - 5, 47, 131, 12 eggs adhere to vegetation eggs adhere to vegetation *similar to silvery minnow to 10 m. thought to occur in open water eggs adhere to vegetation uses nests of 140 other minnows spotfin shiner X X - - low low undersides of fixed objects 5, 39, 47, 131 adhesive eggs sand shiner X X - - low low roots of aquatic vegetation 47, 131 adhesive eggs redfin shiner X - - - - - - 81, 99, 102, 131 utilizes sunfish nests mimic shiner X X X - high high - 47, 93, 131 eggs scattered pugnose minnow* X X - - medium medium - 47, 76, 116 *adult habitat bluntnose minnow X X X - medium medium undersides of fixed objects 47, 93, 131, 136 adhesive eggs fathead minnow X X - - medium medium undersides of fixed objects 6, 47, 120, 136 adhesive eggs blacknose dace X - - - - - - 4, 6, 47, 131 longnose dace X X - - - - - 4, 5, 47, 131 creek chub X - - - - - - 6, 47, 131 fallfish X - - - - - - 47, 123, 131 pearl dace X - - - low low - 6, 131 central stoneroller X - - - - - - 5, 47, 84, 86, 88 striped shiner X - - - low low - 5, 44 black bullhead X X - - high high debris, overhanging banks, rocks 5, 47, 131, 146 yellow bullhead X X - - high high logs, tree roots, boards, debris 5, 47 brown bullhead X X - - medium medium logs, tree roots, boards, debris 7, 47, 136 channel catfish X X X - low low undercut banks, log jams, rocks 47, 131 stonecat X X - - - - nests under boards, rocks, logs 47, 131, 142 tadpole madtom X X - - medium medium under boards, logs, roots, 47, 131, 145 13 cans brindled madtom X X - - medium medium under boards, logs, roots, cans 15, 47, 113 banded killifish X - - - high high - 47, 59 eggs adhere to vegetation burbot X X X c - - - 47, 79, 131 may also spawn in deep water brook stickleback X - - - high high sticks 47, 124, 131, 150, 151 threespine stickleback X - - - low low - 47, 150, 151 ninespine stickleback X X X X high high between rocks 37, 47, 150, 151 fourspine stickleback X - - - high high - 21, 127, 150, 151 trout-perch X X X X low low between rocks 5, 47, 131 white perch X X X - medium medium - 9, 47, 136 white bass X X X X low low - 5, 47, 131, 136 rock bass X X - - low low under rocks, logs 47, 49, 101, 136 green sunfish X X - - high high rocks, logs, clumps of grass 5, 47, 76, 131 pumpkinseed X X - - high high - 47, 67, 136 bluegill X X X - high high - 5, 47 longear sunfish X X X - medium medium - 5, 67, 87 smallmouth bass X X - - low low boulders, docks, logs 5, 47, 97, 129, 131 largemouth bass X X - - medium high - 13, 47, 94, 129, 136, 146 white crappie X X X - medium medium banks, brush, stumps 47, 50, 131 c 14 c favours, but is not restricted to, open areas black crappie X X X - high high - 47, 136, 146 warmouth* X X - - high high stumps, rocks, clumps of vegetation 5, 131 orangespotted sunfish X X - - low low - 5, 47, 76, 100 yellow perch X X X X medium medium rocks, brush, debris 47, 119, 136, 143 sauger X X X - low low - 5, 47, 131 walleye X X X X low low - 47, 52, 105 eastern sand darter* X X X - - - - 56, 69 greenside darter X - - - low low algae-covered rocks 29, 47, 69 147, 148 rainbow darter X - - - low low - 47, 66, 69, 147, 148 Iowa darter X - - - medium medium undercut banks, fibrous roots 47, 69, 147, 148 fantail darter X - - - - - undersides of rocks 47, 69, 147, 148 least darter X - - - high high - 28, 63, 69, 147, 148 johnny darter X - - - - - undersides of rocks, logs 47, 66, 69, 147, 148 logperch X X - - - - - 47, 66, 69, 147, 148 channel darter X X - - - - near large rock 47, 69, 148 blackside darter X - - - - - - 5, 69, 147 river darter* X - - - - - - 27, 69 tesselated darter X - - - - - undersides of objects 46, 69 brook silverside X - - - high high - 42, 47, 75, 15 *hybridizes with pumpkinseed & bluegill *adult habitat *similar to channel and blackside darters eggs adhere to 95 freshwater drum X X X X low low - 5, 47, 131 mottled sculpin X - - - - - crevices, under rocks, burrows 34, 47, 130 slimy sculpin X X X X - - under rocks or logs 47, 133 spoonhead sculpin X X X X - - possibly under rocks or logs 47, 133 deepwater sculpin - - - X - - - 79, 133 rudd* X X - - high high - 14, 76, 125 ruffe X X X c - medium - logs, branches, rocks 48, 104, 134 round goby X X X - low - undersides of logs, cans 65, 66 tubenose goby X X X - medium - undersides of logs, cans 65, 66 16 vegetation eggs pelagic eggs adhesive *hybridizes with golden shiner c spawn in depths of 3m or less Table 2 Strength of association with substrate types and the importance of lake (versus stream) habitat for spawning Great Lakes fishes. A dash (-) indicates that no information was found to indicate that the species utilizes a particular substrate type. For species marked with an asterisk (*) inferences about relationships have been drawn based on adult habitat or c hybridization with other species. The notation ' ' indicates additional information is given, also in the 'Comments' column. References are listed in Table 1. Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Hard-pan clay lake sturgeon - high high high high high - - high low longnose gar - - - - medium high high - - high spotted gar - - - - medium high high - - high bowfin - - - - - high high - - high alewife - high high high high high high high - high gizzard shad - - - high high high high - - high rainbow trout - - - low high - - - - low Atlantic salmon - - - high high - - - - low brown trout - - medium high high low - - - low arctic charr - - - - high medium - - - medium brook trout - - high high medium low - - medium high c high high high low low - - - high lake whitefish - high high high high medium - low - high cisco - medium high high high high medium medium medium high lake trout c 17 Comments c with scattered cobble, rubble Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Hard-pan clay high high high high high high high high high high - - - - - - high high - high shortnose cisco medium medium medium medium medium high high high - high shortjaw cisco medium medium medium medium medium high medium high medium high pygmy whitefish - - - high high high - - - medium round whitefish - - - high high - - - - high rainbow smelt - medium medium high high high low - - medium northern pike - - - low low high high - - high muskellunge - - - - - medium high high - high grass pickerel - - - - - - high high - high central mudminnow - - - - - - high - - high quillback - - - - medium high high high - high longnose sucker - - - - high high - - - medium white sucker - - - medium high medium - - - medium lake chubsucker - - - - medium high high - - high northern hogsucker - - - - high medium - - - low bloater kiyi 18 Comments eggs adhere to substrate near mouths of Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Comments Hard-pan clay streams bigmouth buffalo - - - medium medium high high - - high spotted sucker - - - high medium medium - - medium low silver redhorse - - high high high - - - - low shorthead redhorse - - - high high high - - - low greater redhorse* - high - - high high - - - low goldfish - - - - medium high high high - high northern redbelly dace - - - - medium high high - - high finescale dace - - - - medium high high - - high lake chub - medium - high high high low - - high common carp - - - medium medium medium high - - high cutlips minnow - - medium high high - - - - low brassy minnow - - - - medium high high - - high eastern silvery minnow - - - - - - high - - high silver chub - - - - high medium - - - medium 19 *adult habitat Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Hard-pan clay hornyhead chub - - high high high medium low - - low golden shiner - - - - - high high - - high pugnose shiner - - - - medium high high - - high emerald shiner - medium medium high high high - - - high bridle shiner - - - - - high high - - high common shiner - - - medium high medium - - - low blackchin shiner - - - - high high high - - high blacknose shiner - - - - medium high - - - high spottail shiner - - medium medium high high - - - high rosyface shiner - - high high high high - - - low spotfin shiner - - - - high high high - - medium sand shiner - - - - high high - - - medium redfin shiner - - - - high high low - - medium mimic shiner - - - - medium high low - - high pugnose minnow* - - - - - medium high high - high bluntnose minnow - medium medium high high medium - - - medium 20 Comments *adult habitat Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Hard-pan clay fathead minnow - - - - medium high high - - medium blacknose dace - - - high high medium - - - low longnose dace - - medium high high medium - - - low creek chub - - - high high high - - - low fallfish - - - high high - - - - low pearl dace - - - - high high - - - low central stoneroller - - - high high high - - - low striped shiner - - - - high high low - - low black bullhead - - low low low high high - - high yellow bullhead - - - - medium high high - - high brown bullhead - - - - - high high high - high channel catfish - - high high high high high high - medium stonecat - high high high medium - - - - medium tadpole madtom - - - - medium high high - - medium brindled madtom - medium medium medium medium medium high - - medium banded killifish - - - high high high medium - - high 21 Comments Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Hard-pan clay burbot - high high high high medium medium - - high brook stickleback - - - - medium high high - - high threespine stickleback - - - - medium high high - - high ninespine stickleback - - medium high high medium medium - - high fourspine stickleback - - medium medium medium high high - - high - - - high high high high - - medium white perch medium medium medium medium medium medium medium medium medium high white bass high high high high high medium low low medium high rock bass - - high high high medium medium medium - high green sunfish - - - high high high medium medium - high pumpkinseed - - - - high high - medium - high bluegill - - - - high high medium - - high longear sunfish - - - - high medium - medium medium high - - high high medium - - - high trout-perch c smallmouth bass c medium 22 Comments c no apparent preference c with overlying Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Comments Hard-pan clay gravel largemouth bass - - - low low high high high - high white crappie - - - - medium low low high high high black crappie - - - - high high high - - high warmouth* - - - medium high high medium - - high orangespotted sunfish - - - - medium high high - - high yellow perch - - - medium high high medium medium - high sauger - medium high high high medium low low - medium high high high high high high - - high medium eastern sand darter - - - - medium high low - - low greenside darter - high high high high - - - - low rainbow darter - - high high high high - - - low Iowa darter - - - high high high high - - medium fantail darter - - high high high high - - - low walleye c 23 *hybridizes with pumpkinseed & bluegill c underlying Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Hard-pan clay least darter - - - - - high high - - medium johnny darter - - medium medium high high medium medium - medium logperch - medium medium high high high - - - medium channel darter - high high high high - - - - low blackside darter - - - - high high - - - low medium medium medium high high - - - - low tesselated darter - - - high high high - - - low brook silverside - - - medium medium high - - - high river darter* c freshwater drum medium medium medium medium medium medium medium medium medium high mottled sculpin - high high high high high - - - medium slimy sculpin - high high high high low low - - medium spoonhead sculpin high high high high low - - - - high deepwater sculpin high high high low - - - high high high - - - - - high high - - high rudd* 24 Comments *similar to channel and blackside darters c eggs scattered randomly *hybridizes with golden Common Name Substrate Lake Affinity Bedrock Boulder Cobble Rubble Gravel Sand Silt Clay Comments Hard-pan clay shiner ruffe - - - - high high medium high high high round goby - - high high high medium - - - high tubenose goby - - high high high medium - - - high 25 Table 3 Season, range of water temperature and reproductive guild for Great Lakes fishes. Ranges given indicate temperatures above and below which a species will not spawn. Reproductive guilds correspond to Balon (1975) as follows: A.1 open substratum spawners = broadcast; A.2 brood hider c = hide; B.2 nest spawners = nest. Other information is indicated with a and details are given in the 'Comments' column. For species marked with an asterisk (*) inferences about relationships have been drawn. n/a = not available. Common Name Season Temperature °C Reproductive Guild lake sturgeon spring 13-21 broadcast longnose gar spring 19-29 broadcast spotted gar spring 21-26 broadcast bowfin spring 16-19 broadcast alewife spring 13-21 broadcast gizzard shad spring 16-25 broadcast rainbow trout spring 1-18 hide Atlantic salmon fall 6.7 hide brown trout fall 7-13 hide arctic charr fall 0.5 hide brook trout fall 2-13 hide lake trout fall 3-14 hide lake whitefish fall 1-12 broadcast cisco fall 2-5 broadcast bloater winter n/a broadcast kiyi fall n/a broadcast shortnose cisco spring 4-5 broadcast shortjaw cisco fall n/a broadcast pygmy whitefish fall n/a broadcast round whitefish fall 2-4 broadcast rainbow smelt spring 1-18 broadcast northern pike spring 2-18 broadcast muskellunge spring 8-18 broadcast c c 26 Comments c peak c redd construction begins Common Name Season Temperature °C Reproductive Guild grass pickerel spring 6-12 broadcast central mudminnow spring 13 (approx) broadcast quillback spring 19-28 broadcast longnose sucker spring 2-15 broadcast white sucker spring 6-23 broadcast lake chubsucker spring n/a broadcast northern hogsucker spring 10-16 broadcast bigmouth buffalo spring 13-26 broadcast spotted sucker spring 12-19 broadcast silver redhorse spring 13 c broadcast shorthead redhorse spring 11-21 broadcast greater redhorse spring 17-19 broadcast goldfish spring 16 c broadcast northern redbelly dace spring 21-27 broadcast finescale dace spring 13-18 lake chub spring common carp c Comments c spawning commences c spawning commences broadcast c above 15 broadcast c spring 17-28 broadcast cutlips minnow spring n/a brassy minnow spring 10-13 broadcast eastern silvery minnow spring 13-21 broadcast silver chub spring 19-23 broadcast hornyhead chub spring 18-26 nest golden shiner spring 20-27 broadcast pugnose shiner spring 21-29 broadcast emerald shiner spring 20-23 broadcast c nest 27 possibly triggered by temperature fluctuation commences Common Name Season Temperature °C Reproductive Guild Comments bridle shiner spring 14-26 broadcast common shiner spring 14-28 hide blackchin shiner spring n/a broadcast blacknose shiner spring n/a broadcast spottail shiner spring 18-22 broadcast rosyface shiner spring 20-29 broadcast spotfin shiner spring 18-29 hide sand shiner spring 21 c broadcast c redfin shiner spring 21 c broadcast c mimic shiner spring n/a broadcast pugnose minnow* spring n/a broadcast bluntnose minnow spring 20-28 nest fathead minnow spring 16-29 nest blacknose dace spring 12-27 hide longnose dace spring 11-24 hide creek chub spring 13-27 hide spring c hide c c fallfish 16 c pearl dace spring 17-18 broadcast central stoneroller spring 12-27 hide c striped shiner spring 15-18 hide black bullhead spring 12-25 nest yellow bullhead spring n/a nest brown bullhead spring 14-29 nest channel catfish spring 18-29 nest stonecat spring c 25 nest tadpole madtom spring n/a nest spring c nest brindled madtom 25 28 commences commences *adult habitat commences commences c commences c commences c commences Common Name Season Temperature °C Reproductive Guild banded killifish spring 21-32 broadcast burbot winter 1-10 broadcast brook stickleback spring 8-19 nest threespine stickleback spring 10-23 nest ninespine stickleback spring 9-17 nest fourspine stickleback spring n/a nest trout-perch spring 4-21 broadcast white perch spring 11-15 broadcast white bass spring 13-26 broadcast rock bass spring 14-24 nest green sunfish spring 20-28 nest pumpkinseed spring 13-29 nest bluegill spring 19-27 nest longear sunfish spring 22-25 nest smallmouth bass spring 13-24 nest largemouth bass spring 14-21 nest white crappie spring 14-23 nest black crappie spring 16-26 nest warmouth* spring 21.5 nest orangespotted sunfish spring 18-32 nest yellow perch spring 7-22 broadcast sauger spring 4-12 broadcast walleye spring 4-12 broadcast eastern sand darter spring 14-24 broadcast c 29 Comments c commences *hybridization with bluegill and pumpkinseed Common Name Season Temperature °C Reproductive Guild Comments greenside darter spring 11-19 hide rainbow darter spring 15 c broadcast Iowa darter spring 12-15 broadcast fantail darter spring 19-24 hide least darter spring n/a johnny darter spring 17-25 logperch spring 10-15 hide channel darter spring 19-22 hide blackside darter spring 16 c hide c river darter* spring n/a hide *similar to blackside and channel darters tesselated darter spring 13-19 nest brook silverside spring 20-23 broadcast freshwater drum spring 19-22 broadcast mottled sculpin spring 6-16 nest slimy sculpin spring 5-10 nest c commences broadcast nest c c c commences commences spoonhead sculpin fall 4.5 nest c deepwater sculpin winter 11 c nest c rudd* spring 20-27 broadcast ruffe spring 6-18 broadcast round goby spring n/a nest tubenose goby spring n/a nest 30 ripe male caught at this temp. larvae collected *hybridizes with golden shiner Table 4 Number of species that spawn within each depth stratum. 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Berkeley and Los Angeles, California. 43 Appendix A Great Lakes Basin Fish Species List CODE Common name S011 S012 S013 S014 S031 S041 S042 S051 S061 S062 S063 S071 S072 S073 S075 S076 S077 S078 S079 S080 S081 S082 S091 S093 S094 S095 S096 S097 S099 S100 S101 S102 S121 S131 S132 S133 S136 S141 S152 S161 S162 S163 S164 S165 S166 S167 S168 S169 American brook lamprey Lampetra appendix Northern brook lamprey Ichthyomyzon fossor Silver lamprey Ichthyomyzon unicuspis Sea lamprey Petromyzon marinus Lake sturgeon Acipenser fulvescens Longnose gar Lepisosteus osseus Spotted gar Lepisosteus oculatus Bowfin Amia calva Alewife Alosa pseudoharengus American shad Alosa sapidissima Gizzard shad Dorosoma cepedianum Pink salmon Oncorhynchus gorbuscha Chum salmon Oncorhynchus keta Coho salmon Oncorhynchus kisutch Chinook salmon Oncorhynchus tshawytscha Rainbow trout Onchorhynchus mykiss Atlantic salmon Salmo salar Brown trout Salmo trutta Arctic charr Salvelinus alpinus Brook trout Salvelinus fontinalis Lake trout Salvelinus namaycush Splake Salvelinus hybrid 080x081 Lake whitefish Coregonus clupeaformis Cisco(lake herring) Coregonus artedi Bloater Coregonus hoyi Deepwater cisco(chub) Coregonus johannae Kiyi Coregonus kiyi Blackfin cisco Coregonus nigripinnis Shortnose cisco Coregonus reighardi Shortjaw cisco Coregonus zenithicus Pygmy whitefish Prosopium coulteri Round whitefish Prosopium cylindraceum Rainbow smelt Osmerus mordax Northern pike Esox lucius Muskellunge Esox masquinongy Grass pickerel Esox americanus vermiculatus Tiger muskellunge Esox hybrid 131x132 Central mudminnow Umbra limi Mooneye Hiodon tergisus Quillback Carpoides cyprinus Longnose sucker Catostomus catostomus White sucker Catostomus commersoni Lake chubsucker Erimyzon sucetta Northern hog sucker Hypentelium nigricans Bigmouth buffalo Ictiobus cyprinellus Spotted sucker Minytrema melanops Silver redhorse Moxostoma anisurum Black redhorse Moxostoma duquesnei 44 Scientific Name S170 S171 S172 S173 S181 S182 S183 S184 S185 S186 S187 S188 S189 S190 S191 S192 S193 S194 S195 S196 S197 S198 S199 S200 S201 S202 S203 S204 S205 S206 S207 S208 S209 S210 S211 S212 S213 S214 S215 S216 S217 S218 S231 S232 S233 S234 S235 S236 S237 S244 S251 S261 S262 S271 S281 S282 S283 S284 Golden redhorse Moxostoma erythrurum Shorthead redhorse Moxostoma macrolepidotum Greater redhorse Moxostoma valenciennesi River redhorse Moxostoma carinatum Goldfish Carassius auratus Northern redbelly dace Phoxinus eos Finescale dace Phoxinus neogaeus Redside dace Clinostomus elongatus Lake chub Couesius plumbeus Common carp Cyprinus carpio Gravel chub Erimystax x-punctata Cutlips minnow Exoglossum maxillingua Brassy minnow Hybognathus hankinsoni Eastern silvery minnow Hybognathus regius Silver chub Macrohybopsis storeriana Hornyhead chub Nocomis biguttatus River chub Nocomis micropogon Golden shiner Notemigonus crysoleucas Pugnose shiner Notropis anogenus Emerald shiner Notropis atherinoides Bridle shiner Notropis bifrenatus Common shiner Luxilus cornutus Blackchin shiner Notropis heterodon Blacknose shiner Notropis heterolepis Spottail shiner Notropis hudsonius Rosyface shiner Notropis rubellus Spotfin shiner Cyprinella spiloptera Sand shiner Notropis stramineus Redfin shiner Lythrurus umbratilis Mimic shiner Notropis volucellus Pugnose minnow Opsopoeodus emiliae Bluntnose minnow Pimephales notatus Fathead minnow Pimephales promelas Blacknose dace Rhinichthys atratulus Longnose dace Rhinichthys cataractae Creek chub Semotilus atromaculatus Fallfish Semotilus corporalis Pearl dace Margariscus margarita Silver shiner Notropis photogenis Stoneroller Campostoma anomalum Striped shiner Luxilus chrysocephalus Ghost shiner Notropis buchanani Black bullhead Ameiurus melas Yellow bullhead Ameiurus natalis Brown bullhead Ameiurus nebulosus Channel catfish Ictalurus punctatus Stonecat Noturus flavus Tadpole madtom Noturus gyrinus Brindled madtom Noturus miurus Northern madtom Noturus stigmosus American eel Anguilla rostrata Banded killifish Fundulus diaphanus Blackstripe topminnow Fundulus notatus Burbot Lota lota Brook stickleback Culaea inconstans Threespine stickleback Gasterosteus aculeatus Ninespine stickleback Pungitius pungitius Fourspine stickleback Apeltes quadracus 45 S291 S301 S302 S311 S312 S313 S314 S315 S316 S317 S318 S319 S323 S324 S331 S332 S334 S335 S336 S337 S338 S339 S340 S341 S342 S343 S344 S345 S346 S361 S371 S381 S382 S383 S384 S601 S220 S355 S366 S367 Trout-perch White perch White bass Rock bass Green sunfish Pumpkinseed Bluegill Longear sunfish Smallmouth bass Largemouth bass White crappie Black crappie Warmouth Orangespotted sunfish Yellow perch Sauger Walleye Eastern sand darter Greenside darter Rainbow darter Iowa darter Fantail darter Least darter Johnny darter Logperch Channel darter Blackside darter River darter Tessellated darter Brook silverside Freshwater drum Mottled sculpin Slimy sculpin Spoonhead sculpin Deepwater sculpin CarpxGoldfish Rudd Ruffe Round goby Tubenose goby 46 Percopsis omiscomaycus Morone americana Morone chrysops Ambloplites rupestris Lepomis cyanellus Lepomis gibbosus Lepomis macrochirus Lepomis megalotis Micropterus dolomieu Micropterus salmoides Pomoxis annularis Pomoxis nigromaculatus Lepomis gulosus Lepomis humilis Perca flavescens Stizostedion canadense Stizostedion vitreum vitreum Ammocrypta pellucida Etheostoma blennioides Etheostoma caeruleum Etheostoma exile Etheostoma flabellare Etheostoma microperca Etheostoma nigrum Percina caprodes Percina copelandi Percina maculata Percina shumardi Etheostoma olmstedi Labidesthes sicculus Aplodinotus grunniens Cottus bairdi Cottus cognatus Cottus ricei Myoxocephalus thompsoni Hybrid 181x186 Scardinius erythrophthalmus Gymnocephalus cernuus Neogobius melanostomus Proterorhinus marmoratus APPENDIX B Riverine spawners Common Name References American brook lamprey 1, 78, 132 northern brook lamprey 1, 78, 92 silver lamprey 1, 78 sea lamprey 1, 47, 78 American shad 77 pink salmon 20, 47, 71 chum salmon 11 coho salmon 47 chinook salmon 18, 47 mooneye 1, 47, 141 black redhorse 10, 47, 72, 108 golden redhorse 23, 47, 72 river redhorse 106 redside dace 86, 113 river chub 26, 47, 83 silver shiner 86 ghost shiner 5, 55 northern madtom 45, 126 blackstripe topminnow 85 47
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