Northeastern Naturalist 174 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 22001144 NORTHEASTERN NATURALIST 2V1(o2l). :2117,4 N–1o9. 12 Temporal Stability Patterns in Fish Species Richness, Diversity, and Evenness in Otter Creek, Vigo County, IN Thomas P. Simon1,*, Charles C. Morris1, and John O. Whitaker, Jr.1 Abstract - We used abundance data from fish-population studies (1962–2010) on Otter Creek at Markle’s Dam, Vigo County, IN, to examine temporal variation in stream-fish assemblages. Species-richness variation showed a declining trend during the 50 year study period with a mean of 49 ± 8.8 species per decade (range = 39–62 species). Cumulative study species richness comprised 76 fish species with a mean of 21.3 ± 6.1 species collected per sampling event. Dominant species, based on relative abundance during the 50 year period, included Cyprinella spiloptera (Spotfin Shiner), Pimephales notatus (Bluntnose Minnow), Hybognathus nuchalis (Mississippi Silvery Minnow), and Luxilus chrysocephalus (Striped Shiner). Relative abundance of Spotfin Shiner showed an increasing trend during the 50-year study period, whereas Bluntnose Minnow relative abundance showed a decreasing trend. Cumulative frequency-distribution of species richness normalized by decade showed no significant difference, with the exception of the 1970s, which showed a steep decline in the number of species. Repeated-measures analysis of variance showed no significant differences in species diversity (H') or evenness (Pielou’s J) over the 5-decade study period. Eleven cool-water species including Chrosomus erythrogaster (Southern Redbelly Dace), Perca flavescens (Yellow Perch), Catostomus commersonii (White Sucker), and Ambloplites rupestris (Rock Bass), and lithophilic spawning species such as Nocomis micropogon (River Chub) and Hybopsis amblops (Bigeye Chub) were extirpated during the study period. Potential changes in sedimentation and thermal gradients may have contributed to these observed extirpations. Introduction Spatially heterogeneous environments offer the potential for spatial segregation of species and provide refugia from harsh physical environmental conditions (Schlosser 1991). Temporal variation in the physical environment can influence growth, survival, immigration, and emigration (Karr and Freemark 1985). Spatial and temporal variation alter species richness indirectly by altering nutrients and organic cycling, production, processes at lower trophic levels, resource and habitat availability, and resource use (Fisher et al. 1982, Poff et al. 1997, Probst et al. 2008). Annual species-richness variation is lower in more-diverse stream assemblages (Franssen et al. 2011). Long-term studies (>30 years) of fish-assemblage structure are few; however, regionally important data sets include large midwestern rivers and the Great Lakes, such as the Illinois River (1957–), Wabash River (1966–), and Lake Michigan (1972–) (Beugly and Pyron 2010, Brunnell et al. 2005, Gammon and Simon 2000, 1Department of Biology, 600 Chestnut Street, Science Building, Indiana State University, Terre Haute, IN 47809. *Corresponding author - thomas.simon@indstate.edu. Manuscript Editor: Glen Mittelhauser Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 175 Koehl and Sparks 2002, Pegg and McClelland 2004, Probst et al. 2008). Trends in stable physical environments show that neither species richness nor diversity change temporally (Greenfield and Bart 2005). Physical environmental disturbances within streams and rivers over the last 40 years include increased sedimentation (Beugly and Pyron 2010), habitat fragmentation (Taylor et al. 2008), decline of current velocities (Poff and Allan 1995, Poff et al. 1997), and nutrient-cycle increases (Bunnell et al. 2005, Pegg and McClelland 2004). The Otter Creek study, which began in the late 1870s, is one of a few continuous long-term studies of fish assemblages in small streams, (Blatchley 1938, Gerking 1945, Jordan 1877). Surveys of the stream documented 63 species between 1885 and 1886 (Hay 1894, Jenkins 1887, Jordan 1890). Whitaker and Wallace (1973) sampled 321 sites in Vigo County, including 60 sites in the Otter Creek watershed. Forty-seven fish species (46.5% of the 101 species collected in the entire study) were collected at Markle’s Dam, a site located on Otter Creek in Markle Mill Park. This single site had the highest species richness recorded in the study. Whitaker (1976) reported 57 species within the Otter Creek Markle’s Dam site (1962–1974) based on his 12-year continuous seining study. The concept of stream stochasticity, i.e., response to unpredictable environmental change, is based on study of Otter Creek at Markle’s Dam (Grossman et al. 1982, 1985). Using data collected over 12 years at the site, Grossman et al. (1982) conducted a trait analysis of the 10 most common fish species to rank species by abundance and trophic group, and concluded that the fish assemblage at Otter Creek at Markle’s Dam was determined by stochastic factors; however, these conclusions were not unanimous (Herbold 1984, Rahel et al. 1984, Yant et al. 1984). The current study evaluates temporal changes in species richness, species diversity, and evenness at Markle’s Dam. Our objectives were to assess temporal patterns and identify temporal shifts in fish assemblages over a 5-decade period. Temporal shifts are particularly interesting given the lack of management manipulations at the site, and our results provide some insight into fish-assemblage response over the latter half of the 20th century. Methods Study area and design Otter Creek is a third-order stream draining 326 km2 in Vigo, Clay, and Parke counties in west-central Indiana (Hoggatt 1975). The mainstem of Otter Creek originates near the town of Carbon and flows westward for 38.6 km before entering the Wabash River 4 km North of Terre Haute (Weinman 2007). The Otter Creek watershed is located entirely within the Interior River Lowland ecoregion, an area that is characterized by undulating plains with wide, shallow valleys that were covered by pre-Wisconsian glaciers (Melhorn 1997, Woods et al. 1998). The majority of the land area in the Otter Creek watershed is used for agriculture (41.7%) or is forested (41.4%); residential areas (7.9%) and grass and pastureland (7.6%) account for much of the remaining area (Choi et al. 2005). The qualitative habitat evaluation index (QHEI) for this stream reach ranges from 87–90 QHEI points with a Northeastern Naturalist 176 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 mean of 88.5 ± 0.89 based on 1990–2010 sample events. No habitat measures were completed during the 1960–1989 time period because the QHEI index was not developed until 1989 (Rankin 1989). The Otter Creek Markle Dam reach averages 12.4 m wide and originates 150 m downstream (39.529027oN, 87.346958oW) of the Mill Dam in Markle’s Mill Park (39.527934o N, 87.345928oW), Otter Creek Township (Fig. 1). The Mill Dam is Figure 1. Geographic location of the Markle Dam reach on Otter Creek, Vigo County, IN. Bottom: photograph of Mill Dam in Markle’s Mill Park. Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 177 a national historic landmark originally constructed in 1817, rebuilt in 1938 after burning, and reconstructed in 1984. Limited details of the reconstruction are available, but the dam was dismantled during the mill reconstruction and then rebuilt to the original specifications in 1984. We assume that sediment behind the dam was released during dam removal and that upstream dam pool habitat transitioned into free-flowing run habitat; however, we do not know how long this situation persisted. The dam is a hydrologically stabilizing structure that acts as a sediment trap and mitigates flow downstream and restricts fish passage. We conducted sampling in Otter Creek at the same reach annually (n = 54) during June–August from 1962 to 2010. We grouped data based on annual, 3-yr, 5-yr, and 10-yr periods to evaluate patterns. Because some years had more effort, and to statistically avoid bias of the results by differentially weighting those years that had more effort, we selected 10-yr periods for presentation. Annual analysis increases data resolution, but also increases beta error. We chose time periods for our analysis to balance alpha against beta error; our choice is based on convenience and normal statistical convention. We compiled data into 5 study periods—1962–1970, 1971–1980, 1981–1990, 1991–2000, and 2001–2010. We used a random sample (n = 5) drawn from each decade to perform evaluations and make comparisons. We used cumulative-frequency distribution to determine the number of samples based on distribution of pooled data, to show accumulation of species with increasing number of sample events. To ensure consistency, the same crew leader (J. Whitaker) supervised sampling throughout the entire study period, 1962–2010. Crews conducted sampling during stable flow periods with a minimum of 7 days between events. Researchers avoided sampling immediately after storm events, instead waiting until water clarity and flow rates returned to normal levels, based on the average streamflow that occurred over 7 consecutive days and had a 10-year reoccurrence- interval–period (i.e., 7Q10) discharge mean. We vouchered and deposited fish specimens at the Indiana State University Zoology Collection, Department of Biology, Terre Haute, IN. Field survey methods Each sampling event consisted of 3600 s of seining effort using a 4.5-m seine and a 9-m seine with 6.25-mm standard mesh during daylight hours. We sampled in an upstream direction within a 150-m reach (Whitaker 1976). The reach included a single habitat cycle of riffle, run, and pool habitats. Sampling within the reach included all available instream cover and substrate types. Seining crews included groups of 5 or 6 students. We positioned each seine with a student on each brail. Three to four crew members went upstream a distance equivalent to the width of the seine and then kicked the substrate vigorously to dislodge individual fish by moving in unison in a downstream direction towards the stationary seine. We placed all fish collected into a live-well for later identification. We identified individual fish to species, counted and batch-weighed them by species, and recorded minimum and maximum length for each species. We made our identifications using regional identification manuals (Gerking 1955, Simon 2011, Smith 1973, Trautman 1981). Northeastern Naturalist 178 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 Data analysis We tested for significant differences between decade-long periods using Kruskal Wallis ANOVA, α = 0.05, with a post hoc Tukey’s honest significant difference (HSD) test (Zar 2009). Cumulative frequency distributions included an analysis of decade-delimited fish-collection events in order to compare the number of sample events that maximized the number of species. We standardized these data based on a total of 5 sampling events so that equal numbers of samples were statistically selected to avoid differential weighting of decades (Fig. 2). We measured species α-diversity using the Shannon-Weiner (H') diversity based on the formula, R H' = -Σpi ln pi , i = 1 where pi is the proportion of individuals belonging to the ith species (Shannon 1948). Evenness is a measure of the distribution of individuals over species (Pielou 1975). Pielou’s J is measured based on the formula J = H' , (ln S) where H' is the Shannon-Weiner diversity index and S is the number of species. Figure 2. Standardized cumulative-frequency distribution of species richness from each of 5 decades of sampling at Markle’s Dam on Otter Creek, Vigo County, IN, limited to 5 random observations (1960s = thick black line, 1970s = black dashed line, 1980s = thick gray line, 1990s = gray dashed line and 2000s = thin black line). Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 179 Results During our cumulative study-period sampling (n = 54 events) at the Otter Creek Markle’s Dam site between 1962 and 2010, we collected and identified 203,742 individuals representing 76 species (Appendix 1). The species richness per sample event averaged 21.3 ± 6.1 species (range = 9–33 species). During the 1960s, numerically dominant species included Pimephales notatus (Bluntnose Minnow), Luxilus chrysocephalus (Striped Shiner), and Cyprinella spiloptera (Spotfin Shiner), which comprise 29%, 22%, and 13% of the total decade relative-abundance, respectively. During this decade, we collected 9 species that have not been collected since— Ichthyomyzon unicuspis (Silver Lamprey), Macrhybopsis hyostoma (Shoal Chub), Luxilus cornutus (Common Shiner), Carpiodes velifer (Highfin Carpsucker), Moxostoma macrolepidotum (Shorthead Redhorse), Noturus eleutherus (Mountain Madtom), Pylodictis olivaris (Flathead Catfish), Morone chrysops (White Bass), and Lepomis gulosus (Warmouth). Collections during the 1970s were numerically dominated by Spotfin Shiner and Hybognathus nuchalis (Mississippi Silvery Minnow) comprising 43% and 20% of the total relative abundance for the decade, respectively (Appendix 1). Spotfin Shiner dominance was consistent with the previous decade, while the Mississippi Silvery Minnow increased an order of magnitude compared to the 2% relative abundance it comprised during the 1960s. Bluntnose Minnow and Striped Shiner showed a declining trend in the 1970s collections. We collected 7 species for the first time during the 1971–1979 period: Lepisosteus platostomus (Shortnose Gar), Cyprinus carpio (Common Carp), Erimyzon oblongus (Creek Chubsucker), Moxostoma anisurum (Silver Redhorse), Ictiobus cyprinellus (Bigmouth Buffalo), Lepomis microlophus (Redear Sunfish), and Perca flavescens (Yellow Perch). We failed to collect 8 other native species after the 1970s including River Chub, Notropis boops (Bigeye Shiner), Southern Redbelly Dace, Ameiurus melas (Black Bullhead), Lepomis humilis (Orangespotted Sunfish), Etheostoma spectabile (Orangethroat Darter), Creek Chubsucker, and Yellow Perch. These species are considered cool-water species (Frimpong and Angermeier 2009). Numerically dominant species from 1980 to 1989 included Spotfin Shiner and Gizzard Shad, which comprised 67% and 15% of the total decade relative abundance, respectively. The numerical dominance of Gizzard Shad contrasts with its relative abundance in the 1960s (less than 1%) and 1970s (7%). We collected two species, Amia calva (Bowfin) and Rhinichthys obtusus (Western Blacknose Dace), for the first time during 1980–1989, but did not collect them in subsequent decades. We collected White Sucker, River Carpsucker, and Rock Bass for the last time during the 1980s (Appendix 1). Spotfin Shiner and Gizzard Shad were numerically dominant during 1990–1999, comprising 54% and 16% of the total decade catch, respectively (Appendix 1). We collected Bigeye Chub and Ameiurus natalis (Yellow Bullhead) for the first time during 1990–1999; during this period, Notropis volucellus (Mimic Shiner), Ictiobus niger (Black Buffalo), and Percina sciera (Dusky Darter) returned for the first time since the 1960s. Northeastern Naturalist 180 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 During 2001–2010, Spotfin Shiner dominated collections with Notropis atherinoides (Emerald Shiner), which comprised the second most abundant species. We collected Gambusia affinis (Western Mosquitofish), an invasive species, Cyprinella whipplei (Steelcolor Shiner), and Cottus bairdii (Mottled Sculpin) for the first time during the 2000s. Pimephales vigilax (Bullhead Minnow) returned for the first time since the 1960s. We used our decade-long datasets to compare number of species and number of individuals in each collection and their means and standard deviations. Sampling events from 1962 to 1970 (n = 23) resulted in a total of 173,349 individuals representing 62 fish species (Appendix 1). Species richness per sample event averaged 21 ± 5 species (range = 9–33 species). During 1971–1980 we collected a total of 10,750 individuals (n = 10 sample events; Appendix 1) representing 53 species with an average species richness of 25 ± 2 species (range = 22–29 species) collected during each sampling event. Sampling events from 1981 to 1990 resulted in 14,301 individuals (n = 10 sample events) representing 46 species with an average species richness of 22 ± 4 species collected per event (range - 17–32 species; Appendix 1). During 1991–1999 we collected a total of 2947 individuals (n = 6 sampling events; Appendix 1) representing 45 species with an average species richness of 18 ± 2 species collected per event (range = 16–21). Sampling events from 2001 to 2010 included 2395 individuals (n = 5 sampling events; Appendix 1). We collected a total of 39 species with an average species richness of 18 ± 4 species collected per event (range = 14–24 species). Species richness adjusted for sampling effort by decade (n = 5 sampling events) declined over the five decades of sampling at Markle’s Dam by an average of 5.75 ± 2.99 species per decade (Fig. 3). We observed the maximum net loss during the 1970s (net loss = 9 species), and the smallest species net loss was during the 1990s (net loss = 2 species). We found a significant difference (ANOVA, F = 3.47, P = 0.014) in average number of species collected within year by decade during the 1970s (Tukey HSD = 0.021; Table 1). No other decade showed a significant difference in species richness, H', or J (Table 1). Table 1. Repeated-measures ANOVA with Tukey HSD post hoc test for Shannon-Weiner (H') species diversity and Pielou’s (J) Evenness based on decade temporal change for Otter Creek at Markle’s Dam, Vigo County, IN for 1962 to 2010 (P-values at α = 0.05). H' Evenness Decade 1960s 1970s 1980s 1990s 2000s 1960s 1970s 1980s 1990s 2000s 1960s 1970s 0.9999 0.9994 1980s 0.1277 0.0826 0.1277 0.0826 1990s 0.1504 0.0983 0.9999 0.1054 0.0983 0.9999 2000s 0.1177 0.0758 0.9999 0.9999 0.1177 0.0758 0.9999 0.9999 Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 181 Discussion The structural organization of fish-assemblage response is strongly linked to local habitat variation (Hoeinghaus et al. 2006), which is influenced by hydrologic regime (Poff et al. 1997). Changes in hydrologic-variation patterns are responsible for fish-assemblage structure by introducing variation in disturbance regimes (Poff and Allan 1995). Hydrologically stable sites support specialist species, while hydrologically variable sites support generalist species (Poff and Allan 1995). Hydrologic alterations result in changes in relative abundance. The Otter Creek reach at Markle’s Dam is hydrologically stable as a result of the Mill Dam, Figure 3. Boxand whiskerp l o t s b a s e d on normalized effort (n = 5 samples) showing temporal changes in Otter Creek at Markle’s Dam, Vigo County, IN. A. Shannon-Weiner (H') diversity index, and B. Pielou’s J evenness. Northeastern Naturalist 182 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 and the heterogeneous reach-specific habitat features increase niche complexity and, with a QHEI of 88.5 ± 0.89, this reach has one of the highest habitat scores in southwestern Indiana (T.P. Simon, unpubl data). It is possible that this habitat heterogeneity may have been compromised during the dam reconstruction and events leading up to this repair. Our analysis of 50 years of data from 1962–2010 collections resulted in a cumulative 76 fish species collected from this single site (Appendix 1). Otter Creek at Markle’s Dam species richness represents 75% of the fish fauna known to occur in Vigo County (Whitaker and Wallace 1973) and 38% of the entire fish fauna of Indiana (Simon 2011). The mean species richness within decade by sampling event did not vary significantly (Fig. 2), and normalized effort shows no significant difference in decade cumulative-frequency distribution. The 1960s data is represented by a sigmoidal curve in contrast to the much flatter curves from subsequent decades. In order to detect a decrease in species relative abundance during the 50 years of sampling, we would have to have sampled more frequently to collect the same number of species through time. Our sampling design did not account for seasonality of fish assemblages at this site; some of the species presented in Appendix 1 may be transient and may have been present at the Markle’s Dam location only occasionally for reproduction or refuge and likely would not be collected every event. For example, Dorosoma cepedianum (Gizzard Shad) is a schooling species that can be patchily distributed (Simon 2011). This species was among the most dominant by relative abundance from 1980 to 1999 (Appendix 1). Based on dominant species, loss of native species, and new species occurrences, Markle’s Dam species composition showed a general trend toward declining native species and fewer new native species. Stream fish-assemblage structure and its relation to patterns associated with natural variability need to be understood in order to manage abiotic and biotic conditions in watersheds. Herbold (1984) analyzed Whitaker’s (1976) data and showed similarity in community composition between years; the pool-dwellers, which are composed of bottom-feeding species, showed similarity within years but little similarity between years. Yant et al. (1984) stated that the study site, sampling method, and within-year replications used by Grossman et al. (1982) emphasized season, and their data-analysis methods resulted in more variability than equally plausible alternate choices for those factors. Reanalysis of the data by Yant et al. (1984) concluded that there was actually very little variation in the fish community at this location over the course of 12 years. Gorman (1986) proposed that the presence of Markle’s Dam resulted in more variability in the assemblage compositio n. Changes in species composition over the 5-decade study period show that fish assemblages remained diverse based on measures of species diversity and evenness (Fig. 3); however, species richness and composition of the assemblage were dynamic with decreasing trends over time (Fig. 2). Diversity indices, such as Shannon- Weiner and Pielou’s J, should be regarded as a summary of α-diversity, which is the diversity within a uniform habitat (Whittaker 1972). Both of these measures should Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 183 be considered structural aspects of the assemblage. Spotfin Shiner was among the dominant 3 species during all 5 decade-long periods, but exhibited the highest relative abundance during the period from 1970 to 2010. Many long-term data sets do not include abiotic information for covariate analysis; however, using data from Frimpong and Angermeier (2009), we completed a trait analysis based on temperature and habitat factors such as sedimentation. Eleven species were extirpated during the study period. These species represent cool-water species like Southern Redbelly Dace, Yellow Perch, White Sucker, and Rock Bass, and lithophilic spawning species such as River chub and Bigeye Chub (Appendix 1; Frimpong and Angermeier 2009). Community shifts possibly linked to sedimentation and thermal gradients may have contributed to these observed changes. Our hypothesis is that during the reconstruction of the dam in 1984, sediment that accumulated behind the dam was released downstream. In addition, the river probably returned to shallow, unrestricted flow regimes resulting in warmer water in the reach while the dam was being rebuilt. Substrates downstream may have been blanketed with fine sediments during the removal of the dam. After fish thermally classified as cool-water species were eliminated from the reach and lithophilic spawning minnow species were impacted by the elimination of downstream coarse substrates, neither guild could return. We do not believe that density-dependent predator-prey biotic interactions or other biotic factors directly contributed to changes at Markle Dam because predator numbers declined during this study period. It is important to recognize that factors other than sedimentation and water temperature might explain changes in species over the 50-year study period. As is common in long-term studies that lack information, precise determination of cause and effect is not possible; however, trait analysis may provide some hypotheses for species change that require further evaluation (Frimpong and Angermeier 2009). Indirect effects altering nutrients and organic cycling, production, processes at lower trophic levels from both top-down predator and bottom-up cascading trophic level changes, resource and habitat availability during the reconstruction of the Mill dam, and resource use by dominant species might be equally plausible explanations that either separately or cumulatively may explain patterns. Acknowledgments Special thanks to Indiana State University students in the Vertebrate Zoology classes that assisted in field collection over the years. We especially thank D. Sparks, D.C. Wallace, and R. Benda for field assistance during the past 20 years. Chris Winslow and an anonymous reviewer provided constructive comments. Paul D. McMurray provided assistance organizing an early draft of this manuscript. Literature Cited Beugly, J., and M. Pyron. 2010. Temporal and spatial variation in the long-term functional organization of fish assemblages in a lar ge river. Hydrobiologia 654:215–226. Blatchley, W.S. 1938. The Fishes of Indiana: With descriptions, Notes on Habits and Distribution in the State. The Nature Publishing Company, Indianapolis, IN. 121 pp. Northeastern Naturalist 184 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 Brunnell, D.B., C.P. Madenjian, and R.M. Claramunt. 2005. Long-term changes of the Lake Michigan fish community following the reduction of exotic Alewife (Alosa pseudoharengus). Canadian Journal of Fisheries and Aquatic Sciences 63:2434–2446. Choi, J.Y., B. Engel, and L. Theller. 2005. Watershed Delineation. Available online at http:// pasture.ecn.purdue.edu/~watergen/. Accessed 24 February 2010. Fisher, S.G., J.L. Gray, N.B. Grimm, and D.E. Busch. 1982. Temporal succession in a desert stream-ecosystem following flash-flooding. Ecological Monographs 52:93–110. Franssen, N.R., M. Tobler, and K.B. Gido. 2011. Annual variation of community biomass is lower in more diverse stream-fish communities. Oikos 120:582– 590. Frimpong, E.A., and P.L. Angermeier. 2009. Fish traits: A database of ecological and lifehistory traits of freshwater fishes of the United States. Fisher ies 34(10):487–495. Gammon, J.R., and T.P. Simon. 2000. Variation in a great-river index of biotic integrity over a 20-year period. Hydrobiologia 422:291–304. Gerking, S.D. 1945. The distribution of the fishes of Indiana. Investigations of Indiana Lakes and Streams 3:1–137. Gerking, S.D. 1955. Key to the fishes of Indiana. Investigations of Indiana Lakes and Streams 4:49–86. Gorman, O.T. 1986. Assemblage organization of stream fishes: The effect of rivers on adventitious streams. American Naturalist 128:611–616. Greenfield, D.I., and H.L. Bart, Jr. 2005. Long-term fish-community dynamics from a blackwater stream receiving kraft mill effluent between 1973 and 1988. Hydrobiologia 534:81–90. Grossman, G.D., P.B. Moyle, and J.O. Whitaker, Jr. 1982. Stochasticity in structural and functional characteristics of an Indiana stream-fish assemblage: A test of community theory. American Naturalist 120:423–454. Grossman, G.D., M.C. Freeman, P.B. Moyle, and J.O. Whitaker, Jr. 1985. Stochasticity and assemblage organization in an Indiana stream-fish assemblage. American Naturalist 126:275–285. Hay, O.P. 1894. The lampreys and fishes of Indiana. Annual Reports of the Indiana Department of Geology and Natural Resources 19:146–296. Herbold, B. 1984. Structure of an Indiana stream-fish association: Choosing an appropriate model. American Naturalist 124:561–572. Hoggatt, R.E. 1975. Drainage areas of Indiana streams. US Department of the Interior Geological Survey, Water Resources Division, Indianapolis, IN. 231 pp. Hoeinghaus. D.J., K.O. Winemiller, and J.S. Birnbaum. 2006. Local and regional determinants of stream-fish assemblage structure: Inferences based on taxonomic vs. functional groups. Journal of Biogeography 34:324–338. Jenkins, O.P. 1887. List of the fishes collected in Vigo County in 1885 and 1886. Hoosier Naturalist 2:93–96. Jordan, D.S. 1877. On the fishes of northern Indiana. Proceedings of the Academy of Natural Sciences, Philadelphia 29:42–82. Jordan, D.S. 1890. Report of the explorations made during the summer and autumn of 1888, in the Allegheny region of Virginia, North Carolina, and Tennessee, and in western Indiana, with an account of the fishes found in each of the river basins of those regions. Bulletin of the United States Fish Commission 8:97–173. Karr, J.R., and K.E. Freemark. 1985. Disturbance and vertebrates: An integrative perspective. Pp. 153–168, In S.T.A. Pickett and P.S. White (Eds.). Ecology of Natural Disturbance and Patch Dynamics. Academic Press, New York, NY. Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 185 Koehl, T.M., and R.E. Sparks. 2002. Historical patterns of river stage and fish communities as criteria for operations of dams on the Illinois River. River Research Applications 18:3–19. Melhorn, W.N. 1997. Indiana on ice: The late Tertiary and ice-age history of Indiana landscapes. Pp. 15–27, In M.T. Jackson (Ed.). The Natural Heritage of Indiana. Indiana University Press, Bloomington, IN. 482 pp. Pegg, M.A., and M.A. McClelland. 2004. Spatial and temporal patterns in fish communities along the Illinois River. Ecology of Freshwater Fish 13:125–135. Pielou, E.C. 1975. Ecological Diversity. Wiley, New York, NY. 165 p. Poff, N.L., and J.D. Allan. 1995. Functional organization of stream-fish assemblages in relation to hydrological variability. Ecology 76: 606–627. Poff, N.L., J.D. Allan, M.B. Bain, J.R. Karr, L. Prestegaard, B.D. Richter, R.E. Sparks, and J.C. Stromberg. 1997. The natural flow regime: A paradigm for river conservation and restoration. BioScience 47:769–784. Probst, D.L., K.B. Gido, and J.A. Stefferud. 2008. Natural flow regimes, nonnative fishes, and native fish persistence in arid-land river systems. Ecological Applications 18:1236–1252. Rahel, F.J., J.D. Lyons, and P.A. Cochran. 1984. Stochastic or deterministic regulation of assemblage structure? It may depend on how the assemblage is defined. American Naturalist 124:583–589. Rankin, E.T. 1989. The qualitative habitat evaluation index (QHEI): Rationale, methods, and application. Ohio Environmental Protection Agency, Division of Water Quality Planning and Assessments. Columbus, OH. Schlosser, I.T. 1991. Stream-fish ecology: A landscape perspective. BioScience 41:704–712. Shannon, C.E. 1948. A mathematical theory of communication. The Bell System Technical Journal 27:379–423 and 623–656. Simon, T.P. 2011. Fishes of Indiana: A Field Guide. Indiana University Press, Bloomington, IN. Smith, P.W. 1973. The Fishes of Illinois. University of Illinois Press, Champaign-Urbana, IL. 314 pp. Taylor, C.M., D.S. Millican, M.E. Roberts, and W.T. Slack. 2008. Long-term change to fish assemblages and the flow regime in a southeastern US river system after extensive aquatic ecosystem fragmentation. Ecography 31:787–797. Trautman, M.B.1981. Fishes of Ohio. Ohio State University Press, Columbus, OH. Weinman, M.L. 2007. Otter Creek, Vigo County, 2006 Fish Management Report. Fisheries Section, Division of Fish and Wildlife, Indiana Department of Natural Resources, Indianalopis, IN. 32 p. Whitaker, J.O., Jr. 1976. Fish community changes at one Vigo County, Indiana locality over a twelve-year period. Proceedings of the Indiana Academy of Science 85:191–207. Whitaker, J.O., Jr., and D.C. Wallace. 1973. Fishes of Vigo County, Indiana. Proceedings of the Indiana Academy of Science 51:450–464. Whittaker, R.H. 1972. Evolution and measurement of species diversity. Taxon 21: 213–251. Woods, A.J., J.M. Omernik, C.S. Brockman, T.D. Gerber, W.D. Hosteter, and S.H. Azevedo. 1998. Ecoregions of Indiana and Ohio. (Map poster). US Geological Survey, Reston, VA. Yant, P.R., J.R. Karr, and P.L. Angermeier. 1984. Stochasticity in stream-fish communities: An alternative interpretation. American Naturalist 124:573–582. Zar, J.H. 2009. Biostatistical Analysis. 5th Edition. Pearson, Upper Saddle River, NJ. 960 pp. Northeastern Naturalist 186 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 Appendix 1. Species composition, including minimum and maximum relative abundance and percent relative abundance, over 50 years (1960–2010) segregated into five distinct decade periods for Otter Creek at Markle’s Dam, Vigo County, IN. Number after year is the number of sample events for each decade period. Number of Individuals by Species 1960s (n = 23) 1970s (n = 10) 1980s (n = 10) 1990s (n = 6) 2000s (n = 5) Scientific name/ common name min max %rel min max %rel min max %rel min max %rel min max %rel Ichthyomyzon unicuspis Hubbs and Trautman 1 1 Silver Lamprey Lepisosteus platostomus Rafinesque 1 1 15 15 3 3 Shortnose Gar Amia calva L. 1 1 Bowfin Dorosoma cepedianum (Lesueur) 1 15 6 300 7% 1 1000 15% 10 200 16% 1 1 Gizzard Shad Esox americanus vermiculatus Gmelin 1 1 1 4 1 2 1 1 Grass Pickerel Campostoma anomalum (Rafinesque) 1 570 8% 2 35 1% 1 35 1% 1 100 6% 1 15 1% Central Stoneroller Chrosomus erythrogaster (Rafinesque) 1 1 2 2 Southern Redbelly Dace Cyprinella spiloptera (Cope) 1 576 13% 40 3000 43% 150 3000 67% 30 500 54% 87 500 56% Spotfin Shiner Cyprinella whipplei Girard 4 4 Steelcolor Shiner Cyprinus carpio L. 1 1 1 2 5 5 Common Carp Hybognathus nuchalis Agassiz 1 70 2% 1 1000 20% 2 350 3% 1 100 4% 2 100 5% Mississippi Silvery Minnow Hybopsis amblops (Rafinesque) 1 1 Bigeye Chub Luxilus chrysocephalus Rafinesque 1 9 1 50 1% 1 50 1% 2 2 1 7 Striped Shiner Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 187 Number of Individuals by Species 1960s (n = 23) 1970s (n = 10) 1980s (n = 10) 1990s (n = 6) 2000s (n = 5) Scientific name/ common name min max %rel min max %rel min max %rel min max %rel min max %rel Luxilus cornutus (Mitchill) 3 1693 22% Common Shiner Lythrurus umbratilis (Girard) 2 55 2% 1 200 3% 1 15 5 5 1 20 1% Redfin Shiner Macrhybopsis storeriana (Kirtland) 16 16 3 3 2 2 Silver Chub Macrhybopsis hyostoma (Gilbert) 20 20 Shoal Chub Nocomis micropogon (Cope) 2 4 1 1 River Chub Notemigonus crysoleucas (Mitchill) 1 5 1 12 7 8 3 3 Golden Shiner Notropis atherinoides Rafinesque 1 253 7% 1 200 3% 1 100 2% 10 10 8 132 12% Emerald Shiner Notropis blennius (Girard) 1 11 3 6 4 4 River Shiner Notropis boops Gilbert 1 1 2 2 Bigeye Shiner Notropis buccatus (Cope) 1 87 3% 1 80 2% 1 15 1 25 1% 1 40 5% Silverjaw Minnow Notropis rubellus (Agassiz) 1 8 1 4 1 4 1 14 1% Rosyface Shiner Notropis stramineus (Cope) 1 34 1% 1 25 1% 1 6 15 15 1% 3 50 2% Sand Shiner Notropis volucellus (Cope) 1 3 2 2 1 1 Mimic Shiner Phenacobius mirabilis (Girard) 1 190 2% 1 15 1 10 1 8 1% Suckermouth Minnow Pimephales notatus (Rafinesque) 7 3317 29% 1 250 6% 3 150 2% 3 100 5% 3 20 2% Bluntnose Minnow Northeastern Naturalist 188 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 Number of Individuals by Species 1960s (n = 23) 1970s (n = 10) 1980s (n = 10) 1990s (n = 6) 2000s (n = 5) Scientific name/ common name min max %rel min max %rel min max %rel min max %rel min max %rel Pimephales vigilax (Baird and Girard) 9 9 1 1 Bullhead Minnow Rhinichthys obtusus Agassiz 1 1 Western Blacknose Dace Semotilus atromaculatus (Mitchill) 2 43 2% 2 15 3 5 10 10 1 15 1% Creek Chub Catostomus commersonii (Lacepède) 1 2 1 20 1 2 White Sucker Carpiodes cyprinus (Lesueur) 2 3 1 1 12 12 2 11 1% Quillback Carpiodes carpio (Rafinesque) 19 19 1 2 River Carpsucker Carpiodes velifer (Rafinesque) 20 20 Highfin Carpsucker Erimyzon oblongus (Mitchill) 1 1 Creek Chubsucker Hypentilium nigricans (Lesueur) 1 60 1% 3 40 2% 3 25 1% 1 5 1% 1 4 Northern Hogsucker Ictiobus cyprinellus (Valenciennes in Cuvier 1 1 1 5 1 2 and Valenciennes) Bigmouth Buffalo Ictiobus niger (Rafinesque) 1 1 10 10 Black Buffalo Minytrema melanops (Rafinesque) 1 1 1 1 2 2 Spotted Sucker Moxostoma anisurum (Rafinesque) 1 1 1 4 3 3 2 7 Silver Redhorse Moxostoma duquesnii (Lesueur) 1 1 1 1 1 1 2 2 Black Redhorse Moxostoma erythrurum (Rafinesque) 1 12 1 20 1% 1 50 1% 1 5 1 2 Golden Redhorse Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 189 Number of Individuals by Species 1960s (n = 23) 1970s (n = 10) 1980s (n = 10) 1990s (n = 6) 2000s (n = 5) Scientific name/ common name min max %rel min max %rel min max %rel min max %rel min max %rel Moxostoma macrolepidotum (Lesueur) 9 9 Shorthead Redhorse Ameiurus melas (Rafinesque) 1 1 1 2 Black Bullhead Ameiurus natalis (Lesueur) 4 4 Yellow Bullhead Ictalurus punctatus (Rafinesque) 5 5 1 4 1 1 Channel Catfish Noturus eleutherus Jordan 1 1 Mountain Madtom Noturus miurus Jordan 1 16 1 100 3% 1 2 1 10 1 8 Brindled Madtom Pylodictis olivaris (Rafinesque) 3 3 Flathead Catfish Fundulus notatus (Rafinesque) 1 15 2 4 1 4 1 10 Blackstripe Topminnow Gambusia affinis (Baird and Girard) 1 4 Western Mosquitofish Labidesthes sicculus (Cope) 2 5 2 9 2 100 2% 1 100 4% 5 117 6% Brook Silverside Cottus bairdii Girard 2 2 Mottled Sculpin Morone chrysops (Rafinesque) 6 6 White Bass Ambloplites rupestris (Rafinesque) 1 1 1 1 1 1 Rock Bass Lepomis cyanellus Rafinesque 1 20 1 1 1 1 Green Sunfish Lepomis gulosus (Cuvier in Cuvier and 1 1 Valenciennes) Warmouth Northeastern Naturalist 190 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 Vol. 21, No. 2 Number of Individuals by Species 1960s (n = 23) 1970s (n = 10) 1980s (n = 10) 1990s (n = 6) 2000s (n = 5) Scientific name/ common name min max %rel min max %rel min max %rel min max %rel min max %rel Lepomis humilis (Girard) 1 2 1 1 Orangespotted Sunfish Lepomis macrochirus Rafinesque 1 25 1% 2 10 1 75 1% 1 10 1% 1 15 1% Bluegill Lepomis megalotis (Rafinesque) 1 5 3 3 1 3 1 1 1 1 Longear Sunfish Lepomis microlophus (Günther) 1 5 1 2 3 8 1 1 Redear Sunfish Micropterus dolomieu Lacepède 1 2 1 3 1 6 5 5 1 1 Smallmouth Bass Micropterus punctulatus (Rafinesque) 1 1 1 2 3 4 2 4 1 1 Spotted Bass Micropterus salmoides (Lacepède) 1 3 1 6 1 10 1 1 1 1 Largemouth Bass Pomoxis annularis Rafinesque 20 20 8 8 1 8 1 2 White Crappie Pomoxis nigromaculatus (Lesueur in Cuvier 1 1 1 3 2 10 1 5 1 2 and Valenciennes) Black Crappie Etheostoma spectabile (Agassiz) 1 22 2 2 1 1 Orangethroat Darter Etheostoma blennioides Rafinesque 1 50 2% 6 50 2% 1 40 1% 1 15 1% 4 26 2% Greenside Darter Etheostoma caeruleum Storer 1 50 2% 3 20 1% 3 20 4 20 1% 6 18 1% Rainbow Darter Etheostoma flabellare Rafinesque 1 19 1 10 1 2 2 10 5 5 Fantail Darter Etheostoma nigrum Rafinesque 1 40 1% 1 75 2% 1 15 4 10 1 4 Johnny Darter Perca flavescens (Mitchill) 1 1 Yellow Perch Northeastern Naturalist Vol. 21, No. 2 T.P. Simon, C.C. Morris, and J.O. Whitaker, Jr. 2014 191 Number of Individuals by Species 1960s (n = 23) 1970s (n = 10) 1980s (n = 10) 1990s (n = 6) 2000s (n = 5) Scientific name/ common name min max %rel min max %rel min max %rel min max %rel min max %rel Percina caprodes (Rafinesque) 6 6 1 1 1 1 1 4 Logperch Percina maculata (Girard) 1 5 1 25 1 3 1 3 Blackside Darter Percina sciera (Swain) 10 10 1 1 Dusky Darter