2009 NORTHEASTERN NATURALIST 16(4):607–620 Fish Assemblage Connectivity in the Monongahela River Basin David G. Argent1,* and William G. Kimmel1 Abstract - Tributary and mainstem corridors represent important fish-connectivity avenues in large riverscapes. We evaluated the connectivity of 40 Monongahela River tributaries in southwestern Pennsylvania and their respective mainstem junctions using a variety of gears. Twelve tributaries were so fragmented by physical and water-quality impediments, comparisons could not be made. Among the 28 remaining tributaries, classified as adventitious (1st–3rd order) or ordered (4th–5th order), we evaluated fish communities using the Jaccard coefficient of similarity, a cluster analysis, and a Venn diagram. Adventitious tributaries shared 82% of their total faunal complement with ordered tributaries and 29% with the mainstem, while 70% of the ordered ichthyofauna was common to the mainstem. The ichthyofauna of the adventitious tributary network was more distinct and isolated from the mainstem than that of ordered tributaries. In fragmented riverscapes such as this, islands (tributaries) of biodiversity may warrant special protection. Introduction Contemporary fish species diversity patterns result from a combination of natural and anthropogenic processes occurring at temporal and spatial scales, which in turn influence local distribution patterns (Hocutt and Wiley 1986, Imhof et al. 1996, Ricklefs 1987, Tonn 1990). With respect to fishes, spatial scale can best be described using a hierarchical analysis of watersheds and biotic systems, which tend to be nested and integrated through their connectivity (Frissell et al. 1986, Imhoff et al. 1996). The “River Contimuum Concept” (RCC) provides a framework for understanding the spatial relationships among tributaries that structure fish communities (Vannote et al. 1980). The spatial and temporal distribution of fish communities reflects changes in temperature regimes and habitat types throughout the continuum. Linking the entire system is the connectivity expressed among the mainstem and its network of tributaries, some of which include adventitious streams that do not follow a predictable ordered pattern (Schaefer and Kerfoot 2004, Strahler 1952). Adventitious streams have been defined by various authors (Gorman 1986, Minshall et al. 1985, Schaefer and Kerfoot 2004, Vannote et al. 1980) as usually 1st–3rd order waterways that join a mainstem at least three orders greater in magnitude. Ordered streams follow the hierarchical network as described by Strahler (1952). It has been hypothesized that this large change in stream order can result in abrupt community differences at the interface 1California University of Pennsylvania, Department of Biological and Environmental Sciences, 250 University Avenue, California, PA 15419. *Corresponding author - argent@cup.edu. 608 Northeastern Naturalist Vol. 16, No. 4 point and an overall break in the ordered pattern of the “river continuum” (Schaefer and Kerfoot 2004, Vannote et al. 1980). Connectivity provides linkages facilitating movement of fishes for purposes of feeding, reproduction, and colonization as well as separation of life-history stages. Historically, the ichthyofaunal distribution patterns of large river basins were a reflection of geologic events. Fragmentation or elimination of pathways within these networks compromised the ecological integrity of the entire ecosystem by isolating faunal associations. More recently, anthropogenic factors have played an increasing role as determinants of fish distribution, through alteration of connectivity corridors by dam construction, channelization, diversion, and pollution. Connectivity corridors may differ among riverine systems, particularly in large rivers that receive adventitious tributaries and/or those following an ordered pattern. The Monongahela River basin in southwestern Pennsylvania is an example of a system which includes both adventitious and ordered tributaries. While lotic connectivity within the mainstem has been compromised by the installation of a lock-and-dam network, the majority of mainstem fishes are cosmopolitan among the pools (Argent et al. 2007). Recent surveys of the Monongahela River basin tributaries have documented considerable waterquality degradation due to active and abandoned coal mines, agriculture, and industrial and municipal discharges, and various types of blockages (Argent and Kimmel 2006, Argent et al. 2007, Kimmel and Argent 2006a). As a result, fish communities show varying degrees of fragmentation and resulting loss of connectivity. Our objective was to evaluate patterns of fish-community connectivity among the Monongahela River mainstem and its adventitious and ordered tributaries. We hypothesize (1) that the ichthyofaunas of the adventitious tributaries will differ from those of the ordered tributaries and those of the mainstem and (2) that the tributary benthic assemblages will be isolated from those of the mainstem. Methods Description of the study area The Monongahela River arises from the confluence of the West Fork River and Tygart River at Fairmont, WV and flows 206 km north to Pittsburgh, PA, where it joins the Allegheny River to form the Ohio River (Fig. 1). The 130-km 7th-order reach in Pennsylvania is divided into a series of impoundments by six navigational lock-and-dam structures maintained by the US Army Corps of Engineers. Along its course, the Monongahela River traverses varying land uses including active and abandoned coal mines, the industrialized Mid-Mon Valley, agricultural areas, forested patches, and small towns. Sampling design We employed a variety of sampling strategies to assess fish community composition and targeted faunal associations in the tributaries and 2009 D.G. Argent and W.G. Kimmel 609 mainstem. For logistical reasons, principally scope of sampling effort along with varying weather and flow regimes (i.e., wadeable stream vs. non-wadeable river), fieldwork was conducted during summer and early fall over a four-year period. During the summers of 2003 and 2004, we sampled 40 of the 51 named flowing 1st- to 5th-order warm-water tributaries of the Monongahela River in Pennsylvania (Kimmel and Argent Figure 1. Monongahela River basin of Pennsylvania. The legend indicates stream location by map id number, stream name, and stream classification. No. Name Classification No. Name Classification 1 Streets Run Adventitious 15 Dunlap Creek Ordinate 2 Turtle Creek Ordinate 16 Kelley Run Adventitious 3 Sandy Creek Adventitious 17 Meadow Run Adventitious 4 Peters Creek Ordinate 18 Barneys Run Adventitious 5 Fallen Timber Run Adventitious 19 Fishpot Run Adventitious 6 Lobbs Run Adventitious 20 Tenmile Creek Ordinate 7 Bunola Run Adventitious 21 Rush Run Adventitious 8 Mingo Creek Adventitious 22 Neel Run Adventitious 9 Pigeon Creek Ordinate 23 Muddy Creek Ordinate 10 Sunfish Run Adventitious 24 Wallace Run Adventitious 11 Pike Run Ordinate 25 Middle Run Adventitious 12 Little Redstone Creek Ordinate 26 Whiteley Creek Ordinate 13 Maple Creek Adventitious 27 Georges Creek Ordinate 14 Redstone Creek Ordinate 28 Dunkard Creek Ordinate 610 Northeastern Naturalist Vol. 16, No. 4 2005). The Youghiogheny and Cheat Rivers, 6th-order tributaries, were excluded due to their size. We utilized two-pass backpack electrofishing over wadeable 200-m sampling reaches employing single and multiple units, a methodology shown to be effective in estimating species richness in streams of varying dimensions (Kimmel and Argent 2006b). If no fish were collected over the first 100 m during the first pass, sampling was terminated (Kimmel and Argent 2006b). Of the 40 streams, eight could not be surveyed over the prescribed 200-m reach because they contained barriers to fish passage, and four were contaminated (acid mine drainage and/or sewage) to a degree that precluded the maintenance of viable fish populations. The mouths of five ordered tributaries—Georges, Dunkard, Tenmile, Redstone and Turtle creeks—were surveyed over the prescribed 200-m reach by both boat and backpack electrofishing at each point of connectivity with the mainstem. Large specimens (>250 mm TL) and gamefish were identified in the field and released. All others were fixed in 10% formalin and identified in the laboratory. Using multi-mesh gillnets (38 m long, 2.4 m deep, containing variable panels in 7.62-m lengths: 12.7-cm bar, 10.16-cm bar, 7.62-cm bar, 5.08- cm bar, and 2.54-cm bar mesh) during the summer of 2005, we sampled the ichthyofauna at the points of mainstream junction of the 28 tributaries previously surveyed by electrofishing. This methodology has proven effective in assessing large-bodied (>250 mm TL) riverine fish diversity (Argent and Kimmel 2005). We fished a single net downstream of the confluence of each small tributary surveyed by backpack electrofishing and placed nets both upstream and downstream of the ordered tributary mouths surveyed by both backpack and boat electrofishing. Nets were fished for approximately 24 hours at each site, and captured fish were identified in the field and released. In summer 2006, we surveyed the benthic fishes of the Monongahela River mainstem at the mouth of each of the 28 electrofished tributaries using a trawl described by Herzog et al. (2005). At each site, a two-minute trawl was conducted in the mainstem across each tributary mouth. All captured fish were preserved and returned to the lab for identification and enumeration. Data analysis Each sampled tributary was classified as either adventitious (1st–3rd order) or ordered (4th–5th order) with respect to its mainstem connectivity. We determined fish species richness and relative abundance for each tributary and mainstem confluence point by pooling data collected (e.g., backpack and/or boat electrofishing for each tributary and gillnetting and benthic trawling on the mainstem) at each sampling site. To evaluate the ichthyofaunal similarity among tributaries within the Monongahela River basin, we used the average linkage euclidean distance clustering procedure (Digby and Kempton 1994, Krebs 1989, Minitab 1996). The resulting dendogram grouped streams with similar fish species composition. We used these groupings to further evaluate the fish community similarity between adventitious and ordered tributaries. 2009 D.G. Argent and W.G. Kimmel 611 We compared fish communities at each tributary/mainstem junction with the Jaccard coefficient of community similarity (JCS; Jaccard 1901, Krebs 1989): JCS = a / (a + b + c), where a = number of species found in both the tributary and the river (tributary mouth), b = number of species found only in the river (tributary mouth), and c = number of species found only in the tributary. Jaccard coefficient values were then averaged among adventitious and ordered categories for comparison. We constructed Venn diagrams (Venn 1880) to evaluate species composition relationships among adventitious and ordered tributaries and mainstem locations. We targeted two benthic fish communities, the buffalo/redhorse/carpsucker complex (BRC; Argent et al. 2007) and the darter species association to further evaluate tributary-mainstem connectivity. The BRC was treated as a complex since it represents an integral component of the northeastern United States’ “big river” ichthyofauna (Pflieger 1971) and includes a number of species recognized as “species of special concern” in Pennsylvania (Argent et al. 2007). Results and Discussion Of the 28 tributaries sampled during this survey, 16 were identified as adventitious and 12 as ordered. A total of 2406 fish representing four families and 29 species/hybrids were collected from adventitious tributaries, while 4311 fish representing nine families and 51 species/hybrids came from ordered branches. A total of 93 BRC individuals were captured from eight of the 12 ordered streams, while adventitious streams harbored no members of this complex. Darter captures totaled 1588 and 534 in ordered and adventitious streams, respectively. Composite samples of gill-netting and trawling at mainstem confluence sites revealed 999 individuals representing 10 families and 33 species/hybrids from adventitious streams. Nearly 50% of the individuals collected from these streams were of one species, Notropis volucellus Cope (Mimic Shiner). Ordered tributary confluences yielded 451 individuals representing nine families and 32 species/hybrids. Sixteen individuals which could only be identified as Notropis species were not included in our analyses. Ninety-three BRC individuals were captured from the mainstem. A comprehensive list of the entire ichthyofauna described in this study is given in Argent et al. (2007). In general, species richness among Monongahela River tributaries increased from adventitious through ordered branches, resulting in two distinct categories. Clustering relationships based on species richness grouped seven adventitious and nine ordered streams (Fig. 2). The grouping of adventitious streams possessed species richness values of seven or less. Species richness among ordered tributaries ranged from 17 in Dunlap Creek to 30 and 37 in Redstone and Tenmile Creeks, respectively. The high degree of community 612 Northeastern Naturalist Vol. 16, No. 4 similarity (Fig. 2) between Redstone and Tenmile Creeks may be a result of their geographic proximity (Fig. 1). These tributaries may be considered reservoirs of diversity for the basin because 60 and 73% of tributary (ordered and adventitious) species richness is contained in Redstone and Tenmile Creeks, respectively (Fig. 3). Figure 2. Similarity dendogram of adventitious and ordered (denoted by *) streams grouped by the average linkage Euclidean distance clustering procedure. Figure 3. Proportions of Monongahela River tributary species richness. 2009 D.G. Argent and W.G. Kimmel 613 Deviations from the general clustering pattern among the tributaries are best explained by breaks in connectivity attributable to anthropogenic influences. Three ordered streams (Little Redstone, Peters, and Pigeon creeks) that are impaired by various cultural stressors clustered with one or more adventitious tributaries (Fig. 2). Little Redstone Creek was most similar to Sunfish Run, a 1st-order adventitious tributary which yielded ten species. Historically, Little Redstone Creek was impacted by acid mine drainage discharges from surface coal mining activity, but the current water quality is best described as net alkaline. Although impacted by iron precipitates, Little Redstone Creek harbors 11 species, eight of which were shared with Sunfish Run. Peters Creek paired with 2nd-order Meadow and Streets runs, both impacted by sewage and neither in close proximity to each other nor to Peters Creek. Our sampling station on Peters Creek was downstream of a consolidated sewage overflow and discharges from the Clairton Municipal Authority and US Steel Corporation’s Clairton Works. As a result, Peters Creek receives both continuous treated and intermittent untreated discharges. Field measurements at the time of sampling yielded total dissolved solids (TDS) values in excess of 700 mg/l, nearly double the 400mg/l recommended by Black (1977) for the maintenance of a diverse fish population. Fourth-order Pigeon Creek clustered with 2nd-order Maple Creek, which joins the mainstem 16 km downstream. The Pigeon Creek sampling station was located approximately 2 km downstream from an industrial facility that produces insoluable sulfur products and 3 km downstream from an acid mine drainage treatment facility. These combined discharges yielded stream TDS values of 1235 mg/l, triple the limit recommended by Black (1977), and supported lower fish diversity and abundance than expected for a stream of this size. Mingo Creek, a 3rd-order adventitious tributary paried with ordered Pike Run. Both watersheds, while culturally impacted to varying degrees, maintain significant forested areas that protect their riparian corridors. Mingo Creek flows through a county park that protects for public use 1050 ha of the watershed, including 5 km or nearly half of the stream corridor. Nineteen of the 24 species/hybrids collected in Pike Run were also common to Mingo Creek. Connectivity among the tributaries and Monongahela River mainstem is strongest among the ordered tributaries as shown by the mean JCS values (Fig. 4) and Venn diagram (Fig. 5). Jaccard coefficient values ranged from 0.05 to 0.34 (Tenmile Creek) among ordered streams and from 0 to 0.10 among adventitious streams (Fig. 4). Adventitious tributaries share 82% of their total faunal complement with ordered tributaries and 29% with the mainstem, while 70% of the ordered ichthyofauna is common to the mainstem (Fig. 5). There were six and 12 species exclusive to ordered and mainstem sites, respectively, while no species were exclusive to adventitious tributaries (Fig. 5). 614 Northeastern Naturalist Vol. 16, No. 4 Adventitious tributaries appear to have limited linkage to the mainstem as evidenced by the relatively low JCS values (Fig. 4) among paired tributary and mainstem sites, and by their faunal associations, which contain few riverine species (Fig. 5). The disconnect between the adventitious tributaries and the mainstem can be further illustrated by the BRC, none of which were collected at adventitious/mainstem confluences (Fig. 5). During low-flow conditions, these tributaries may be physically separated from the mainstem and therefore isolated from members of this complex. By contrast, the ordered/ mainstem linkage is frequented by members of the BRC at points of confluence (Fig. 5). However, since neither the BRC fishes nor other members of the Monongahela River ichthyofauna have been shown to be obligate tributary spawners (Cooper 1983), their presence may signify resident populations or occasional usage for purposes of life-stage separation, feeding, or possibly occupation of refugia. Also, high flows of the river mainstem often flood the mouths of large tributaries, presumably facilitating the mixing of members of their respective faunas at such times. Darter assemblages of the adventitious and ordered tributaries exhibited a disconnect with that of the mainstem and to a lesser degree with each other, presumably due to limited mobility and habitat preference (Kuehne and Barbour 1983). While mainstem and tributary junctions do share some species (Fig. 5), their proportional abundance (Fig. 6) indicates that the tributary Figure 4. Mean (+ S.D.) Jaccard coefficient of similarity for adventitious and ordered streams. Figure 6 (opposite page bottom). Proportional abundance of darters collected among adventitious and ordered tributaries and mainstem confluence sites. 2009 D.G. Argent and W.G. Kimmel 615 Figure 5. Venn diagram depicting those species collected among adventitious and ordered tributaries and mainstem confluence sites (see Appendix 1 for scientific names). 616 Northeastern Naturalist Vol. 16, No. 4 darter assemblages are largely isolated from those of the mainstem. Of the six riverine darter species (Fig. 5), two—Percina copelandi Jordan (Channel Darter), an obligate mainstem species, and Etheostoma nigrum Rafinesque (Johnny Darter), a cosmopolitan species—comprised 78% of the total darter complement (Fig. 6). The darter composition among tributary sites also differed (Fig. 6). While both adventitious and ordered streams were dominated by Etheostoma caeruleum Storer (Rainbow Darter) (84% and 64% of total populations, respectively), adventitious streams supported only three species of darter, while six were common to ordered branches (Figs. 5 and 6). The RCC has been expanded to include temporally isolated floodplain waterbodies of major rivers (Amoros and Bornette 2002) and is challenged by the existence of adventitious tributaries (Schaefer and Kerfoot 2004) that do not fit the hierarchical pattern and constitute a break in the continuum. The ichthyofauna of the adventitious tributary networks has been shown to be more distinct and isolated from the mainstem as one proceeds upstream (Schaefer and Kerfoot 2004). The tributary network of the Monongahela River consists of both adventitious and ordered branches, with the ordered branches closely connected to the mainstem fish community while the adventitious streams share few mainstem species. However, usage of ordered tributaries by mainstem fishes may be temporary and transitory (Schaefer and Kerfoot 2004). Biodiversity reservoirs exist in Tenmile and Redstone Creeks, which together comprise over 80% of the total tributary complement and may provide refugia and sources of colonizers. Patterns of connectivity along the Monongahela River and its tributaries are largely fractured by physical blockages to fish passage and/or point and non-point sources of pollutants, as evidenced by the 12 tributaries that could not be sampled. In fragmented riverscapes such as that of the Monongahela River, “islands” of biodiversity may warrant special protection by regulatory agencies. Acknowledgments We thank our research technicians, the US Environmental Protection Agency, and the Pennsylvania Fish and Boat Commission for their assistance with field collections. Funding for this project was provided by the US Fish and Wildlife Service State Wildlife Grant #WM-6-02G-0062, Wildlife Resource Conservation Fund Contract # WRCP-04019, and the Faculty Professional Development Center at California University of Pennsylvania. Two anonymous reviewers are also recognized for their comments which helped to strengthen this paper. Literature Cited Amoros, C., and G. Bornette. 2002. Connectivity and biocomplexity in waterbodies of riverine floodplains. Freshwater Biology 47:761–776. Argent, D.G., and W.G. Kimmel. 2005. Efficiency and selectivity of gill nets for assessing fish community composition of large rivers. 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Venn, J. 1880. On the diagrammatic and mechanical representation of propositions and reasonings. Philosophical Magazine and Journal of Science. Series 5. 2009 D.G. Argent and W.G. Kimmel 619 Appendix 1. Fishes listed in Figure 5 with their associated authority C ommon Name Scientific Name Skipjack Herring Alosa chrysochloris (Rafinesque) Gizzard Shad Dorosoma cepedianum (Lesueur) Longnose Gar Lepisosteus osseus (L.) Mooneye Hiodon tergisus Lesueur Central Stoneroller Campostoma anomalum (Rafinesque) Spotfin Shiner Cyprinella spiloptera (Cope) Common Carp Cyprinus carpio L. Striped Shiner Luxilus chrysocephalus Rafinesque Common Shiner Luxilus cornutus (Mitchill) Silver Chub Macrhybopsis storeriana (Kirtland) River Chub Nocomis micropogon (Cope) Golden Shiner Notemigonus crysoleucas (Mitchill) Emerald Shiner Notropis atherinoides Rafinesque Rosyface Shiner Notropis rubellus (Agassiz) Sand Shiner Notropis stramineus (Cope) Mimic Shiner Notropis volucellus (Cope) Channel Shiner Notropis wickliffiTrautman Bluntnose Minnow Pimephales notatus (Rafinesque) Blacknose Dace Rhinichthys atratulus (Hermann) Creek Chub Semotilus atromaculatus (Mitchill) River Carpsucker Carpiodes carpio (Rafinesque) Quillback Carpiodes cyprinus (Lesueur) Highfin Carpsucker Carpiodes velifer (Rafinesque) White Sucker Catostomus commersonii (Lacepède) Northern Hogsucker Hypentelium nigricans (Lesueur) Smallmouth Buffalo Ictiobus bubalus (Rafinesque) Black Buffalo Ictiobus niger (Rafinesque) Silver Redhorse Moxostoma anisurum (Rafinesque) River Redhorse Moxostoma carinatum (Cope) Black Redhorse Moxostoma duquesnii (Lesueur) Golden Redhorse Moxostoma erythrurum (Rafinesque) Shorthead Redhorse Moxostoma macrolepidotum (Lesueur) Yellow Bullhead Ameiurus natalis (Lesueur) Channel Catfish Ictalurus punctatus (Rafinesque) Stonecat Noturus flavus Rafinesque Flathead Catfish Pylodictis olivaris (Rafinesque) White Bass Morone chrysops (Rafinesque) Hybrid Striped Bass Morone hybrid Rock Bass Ambloplites rupestris (Rafinesque) Green Sunfish Lepomis cyanellus Rafinesque Pumpkinseed Lepomis gibbosus (L.) Bluegill Lepomis macrochirus Rafinesque Sunfish Hybrid Lepomis hybrid Smallmouth Bass Micropterus dolomieu Lacepède Spotted Bass Micropterus punctulatus (Rafinesque) Largemouth Bass Micropterus salmoides (Lacepède) White Crappie Pomoxis annularis Rafinesque 620 Northeastern Naturalist Vol. 16, No. 4 C ommon Name Scientific Name Black Crappie Pomoxis nigromaculatus (Lesueur) Greenside Darter Etheostoma blennioides Rafinesque Rainbow Darter Etheostoma caeruleum Storer Fantail Darter Etheostoma flabellare Rafinesque Johnny Darter Etheostoma nigrum Rafinesque Variegate Darter Etheostoma variatum Kirtland Banded Darter Etheostoma zonale (Cope) Yellow Perch Perca flavescens (Mitchill) Logperch Percina caprodes (Rafinesque) Channel Darter Percina copelandi (Jordan) Sauger Sander canadensis (Griffith & Smith) Walleye Sander vitreus (Mitchill) Saugeye Sander hybrid Freshwater Drum Aplodinotus grunniens Rafinesque Mottled Sculpin Cottus bairdii Girard