August 20, 2008
Habitat Use of Grass Pickerel Esox Americanus Vermiculatus in Indiana Streams
By Cain, Michelle L Lauer, Thomas E; Lau, Jamie K
ABSTRACT. - One hundred and twenty five sites from 91 rivers and streams in Indiana were evaluated from 1990-2005 to determine grass pickerel Esox americanus vermiculatus habitat use. Macro-habitat analysis was conducted using the Qualitative Habitat Evaluation Index (QHEI) and had no significant relationship with grass pickerel catch. However, the individual components that made up the QHEI metric of "pool/glide and riffle/run quality" were significant and showed that grass pickerel avoided riffle habitats. At selected (n = 9) stream sites, microhabitat analysis indicated grass pickerel were always associated with either aquatic macrophytes or logs/woody debris. Although the Index of Biotic Integrity has been used to infer overall stream health and community fish quality, this index was minimally related to grass pickerel catch. The results of this study suggested that grass pickerel preferred habitat types in Indiana streams of aquatic macrophytes, logs/woody debris and slow moving water.INTRODUCTION
Physical and chemical features needed for fish existence, such as suitable water quality, migration routes, spawning grounds, feeding sites, resting sites and shelter from enemies and adverse weather is typically referred toas "habitat" (Orth and White, 1999). Habitat types vary among lake, river and stream environments (e.g., riffle areas) , and individual species may have unique habitat characteristics. For example, stream habitat structures the types of fishes within a stream system (Sullivan et al, 2004; Lau et al, 2006) and thus, proper management of habitat cannot be ignored (Orth and White, 1999). Defining, understanding and measuring relative habitat characteristics may be the key to understanding patterns of community diversity among stream fishes (Gorman and Karr, 1978). In addition, physical habitat characteristics (e.g., water depth, current velocity and substrate) are important factors that influence fish communities (Hynes, 1970; Gorman and Karr, 1978; Schlosser, 1982a, b; Moyle and Vondracek, 1985). When stream habitat is degraded, the associated fish community suffers (Etnier, 1972; Gorman and Karr, 1978; Scarnecchia, 1988; Shields et al, 1998; Wiehert and Rapport, 1998; Walser and Bart, 1999). Determining physical habitat features can provide a greater understanding of the mechanisms structuring the fish community.
Grass pickerel Esox americanus vermiculatus (Crossman, 1966) is the smallest member of the family Esocidae which includes muskellunge Esox masquinongy, northern pike Esox lucius, chain pickerel Esox niger and redfin pickerel Esox americanus americanus (Crossman, 1966). Habitat requirements of these fish have some similarities, but have not been identified completely for the entire family (Foster, 1980; Stegeman, 1989; Pflieger, 1997). For example, muskellunge in Tennessee streams prefer pools with plenty of sunlight but often remained in the shaded edges of the pool (Parsons, 1959), whereas northern pike typically thrive in lakes with adequate flooded vegetation, cool water and large-sized prey (Paukert and Willis, 2003). Grass pickerel habitat is thought to be similar to other esocids, as all members of this family are similar morphologically (Foster, 1980) and functionally as lie-inwait predators (Moyle and Cech, 2004). However, comparatively little research has been conducted on grass pickerel habitat despite the fact that they are common and widely distributed in Midwestern lakes and streams (Crossman, 1966). Their small size (grass pickerel only grow to approximately 350 mm total length; Pflieger, 1997) may determine and define their habitat niche, which may be similar to the young of other sympatric genera (e.g., muskellunge; Murry and Farrell, 2007), but different when larger (e.g., northern pike; Headrick and Carline, 1993). Lasdy, their name suggests some association with vegetation, presumably because of the cover where they are typically found (Scott and Crossman, 1973; Trautman, 1981)
Research on grass pickerel habitat has been primarily limited to three biogeographical ecosystems: lakes in Wisconsin (Klienert and Mraz, 1966), streams and artificial lakes in Oklahoma (Ming, 1968) and ponds in Canada (Crossman, 1962; Foster, 1980). Although these studies can be used to draw parallels with fish found in Indiana, extensive comparisons pose problems. First, streams vary throughout Indiana from natural to anthropogenically altered and in a variety of ecotypes (Homoya et al, 1984). Second, Indiana's landscape is 70% row crop agriculture (Indiana Agricultural Statistics Service, 2000), which can have negative effects on the surrounding stream communities including sedimentation and channel erosion (Berkman and Rabeni, 1987 Gammon, 1995). Third, livestock are commonly allowed in Indiana stream and riparian areas, and degrade habitat quality by stirring up sediment, eroding stream banks and increasing ammonia and nitrate concentrations (Gammon, 1995). In contrast, Wisconsin's land use is 43% agriculture (Wang et al, 1997), whereas in Ontario Canada, agriculture represents only 17% of the land use (Canadian Ministry of Agriculture, 2004) . Although geographic areas like Oklahoma have similar landscape use to Indiana (Oklahoma Agriculture Statistics, 2005), grass pickerel only occur in the southeastern corner of the state (Crossman, 1966). Thus, confounding problems may mask our ability to delineate specific habitat dimensions for grass pickerel as they exist in Indiana, based on related published literature.
Information on specific grass pickerel habitat in the Midwestern U.S. has been alluded to by several investigators. Kwak (1988) identified the use of floodplain areas by grass pickerel in the Kankakee River in Illinois, just east of the Indiana-Illinois border. Although he did suggest flow influences the use of these habitats, no specific habitat quantification of either the ditch or the pool areas associated with them were made. Carline and Klosiewski (1985) found abundant levels of grass pickerel in Ohio and showed that grass pickerel abundance increased with the consu*uction and placement of rock deflectors. However, habitat use and selection by grass pickerel was not indicated. Similarly in another Ohio stream, Trautman and Gartman (1974) found grass pickerel in weedy and static habitats and were negatively affected by siltation and increased flow. Unfortunately, their habitat measurements likely came from field notes or unpublished data and were not rigorously examined in the published study. Stream fish habitat in Michigan was documented by Zorn et al. (2004), but little specific information was found for grass pickerel. Lasdy, Larimore (1961) showed a high abundance of grass pickerel in Jordan Creek, Illinois and suggested diese fish were associated with aquatic vegetation when captured. However, this study focused on capture methodology, with habitat descriptions peripheral to the main objective.
Our objective was to determine whether grass pickerel catches were related to available habitat in streams and rivers throughout Indiana. We examined the relationships between grass pickerel catches and Qualitative Habitat Evaluation Index, Index of Biotic Integrity, QHEI individual components, and water quality variables because habitat and fish community quality have been show to be related in East Central Indiana Streams (Lau et al., 2006). We also examined the relationship between QHEI and IBI. These analyses evaluated whether a composite measure of habitat (QHEI) could be associated with grass pickerel use, or whether individual, specific metrics were needed. In some instances when grass pickerel were captured, the type of habitat sampled was assessed visually, and represented a micro level assessment not possible using QHEI alone. Lastly, we compared grass pickerel catch with IBI scores to determine whether grass pickerel and the quality of the fish community were related.
METHODS AND MATERIALS
One hundred and twenty five stream sites in Indiana with drainage areas ranging from 0.75-750 km^sup 2^ were sampled for grass pickerel from 1990-2005 (Fig. 1). These stream sites included both channelized (anthropogenically altered) and un-channelized (natural) sites from 50 of Indiana's 92 counties; no attempt was made to determine the exact length of time since the alterations occurred. However, recently channelized streams were characterized with no channel sinuosity, little riparian vegetation and little or no in- stream cover. Sites recovering from channelization exhibited little or no channel sinuosity, had woody riparian vegetation as well developed as forested areas and some in-stream cover. Lastly, unaltered sites showed well defined channel sinuosity, large riparian areas and extensive in-stream cover. To reduce the confounding affects of high discharge on measuring stream habitats, streams were typically sampled when discharge levels were below United States Geological Survey summer daily median values (http:// waterdata.usgs.gov/ in/nwis/rt) accessed at the time of sampling.
Fish were collected using tote barge, backpack or boat electrofishing following Index of Biotic Integrity (IBI) methodology for Eastern Corn Belt Plain of Indiana (Simon and Dufour, 1998). The length of stream sampled was a minimum of 50 m for small streams (
Habitat characteristics of sample sites were evaluated using two methods: the Qualitative Habitat Evaluation Index (QHEI) (Rankin, 1989) and visual assessment of microhabitat cover. A QHEI was conducted on every site at the time of fish collection. Gradient metric scores for the QHEI were computed using the United States Geological Survey topographic maps (1:24,000 scale). Microhabitat evaluation was performed at each site where a grass pickerel was found for the May to Aug. 2005 sampling season (n = 9). A visual assessment of cover (type, amount) was made whenever grass pickerel were collected. Proportions of in-stream cover where grass pickerel were found were compared with total in-stream cover based on overall estimates from visual observations performed at the site. Components of the QHEI metric for in-stream cover were evaluated. The cover was classified into one of three microhabitat groups: aquatic macrophytes, logs/woody debris or no cover. These classifications were recorded for each area where grass pickerel were collected from each stream reach.
Grass pickerel were assumed to be associated with some type of habitat. This a priori assumption suggested that we compare grass pickerel catch (response variable) with habitat features (predictor variables) of Indiana streams (e.g., temperature, vegetation, QHEI) where this fish was present. Because QHEI values are positively and linearly related to fish community quality in East Central Indiana streams as measured by the IBI (Sullivan et al, 2004), a comparison of both grass pickerel catch and QHEI was also made to the IBI values. Thus, grass pickerel catches were regressed against total QHEI values using a simple linear regression analysis and against all individual QHEI metrics (Table 1). We also separated the pool/ glide and riffle/run quality QHEI metric into separate pool quality and riffle quality components (Table 1). These individual components were subsequendy compared with grass pickerel catches using a best subsets linear regression analysis to further identify specific components of grass pickerel habitat. The best subsets analysis produces multiple models that compare all possible combinations of predictor variables (e.g., habitat components). Typically, we could identify the "best" model by comparing the adjusted r values generated by each (highest value = best fit; Zar, 1999). A simple linear regression analysis was also used to compare microhabitat cover to grass pickerel catches. Lastly, grass pickerel catch was regressed against average stream width, conducdvity, dissolved oxygen, pH and temperature using a simple linear regression to determine whether these factors influenced grass pickerel habitat use. All analyses were run using Minitab v. 14 software using alpha = 0.05.
A total of 111 species was observed from 107 sites where grass pickerel were observed. The number of fish species at each site ranged from 1 to 41, and all sites used in analysis contained 1 to 25 grass pickerel (Table 2). The species most commonly occurring by number with grass pickerel were green sunfish Lepomis cyanellus, bluntnose minnow Pimephales notatus, johnny darter Etheostoma nigrum, bluegill Lepomis macrochirus, creek chub Semotilus atromaculatus, longear sunfish Lepomis megahtis, white sucker Catostomus commersonii and yellow bullhead Ameiurus natalis.
Total QHEI scores ranged from 14-95 (Table 2), representing a range of habitat quality from poor to excellent following the classification of Rankin (1989). However, there was no linear relationship between grass pickerel catch and QHEI (r^sup 2^ = 0.008, df = 123, P = 0.334), or the six QHEI metrics and grass pickerel catch (r^sup 2^ 0.09 for all six comparisons). However, when the individual components of the pool/ glide and riffle/run quality metric (pool max depth, morphology, velocity, riffle depth, run depth, riffle/run substrate and riffle/ run embeddedness) were compared with grass pickerel catches using a best subsets regression, a relationship was identified (r^sup 2^ = 0.70, df = 16, P = 0.001). In this analysis, riffle depth, morphology, riffle/run substrate and riffle/run embeddedness were negatively related to grass pickerel catches.
Microhabitat analysis (n = 9 sites) indicated grass pickerel were always associated with either aquatic macrophytes (77%) or logs/ woody debris (23%). In addition, when the proportion of these cover types in the stream increased, there was a concomitant increase in grass pickerel catch (r^sup 2^ = 0.57, df = 7, P = 0.03).
Because all sites (n = 125) evaluated contained grass pickerel, we examined the relationships between IBI and QHEI, and grass pickerel catch and QHEI to determine whether habitat factors were structuring the fish community and the catch of grass pickerel. Index of Biotic Integrity and QHEI were related (Fig. 2; Table 3), indicating that fish community quality increased with increasing habitat quality. Most of the variation was explained by the pool quality, instream cover and substrate metrics.
Water quality data was collected at 83 sites. Temperatures ranged from 11-32.3 C, pH ranged from 7.16-9.68, turbidity ranged from 0- 140 NTUs, conductivity ranged from 149-1600 S/cm and dissolved oxygen ranged from 0.25-16 mg/l. Values for all these factors, except temperature were significantly related to grass pickerel catch, but the relationships were biologically weak, with the possible exception of average stream width (Table 4).
Habitat factors have not been extensively evaluated for grass pickerel with studies limited to areas in Wisconsin, Okalahoma and Canada (Klienert and Mraz 1968; Ming, 1968; Foster, 1980). In Wisconsin lakes, grass pickerel used shallow vegetated areas (Klienert and Mraz, 1968). Similarly, in Oklahoma streams, grass pickerel were found in headwaters among dense beds of vegetation and in Canadian ponds grass pickerel were almost always associated with vegetation in shallow water areas (Foster, 1980). By comparison, grass pickerel in Indiana also selected habitat components with in- stream cover of submerged aquatic vegetation. However, our finding of grass pickerel association with logs/woody debris in low flow areas was unique. In fact, grass pickerel were found 100% of the time associated with either aquatic macrophytes or logs/woody debris; these cover types are likely used to aid in the ambush of prey (Moyle and Cech, Jr., 2004). The concept of concealment while feeding is similar to other piscivorous fish such as muskellunge (Hanson and Margenau, 1992), northern pike (Paukert and Willis, 2003) and largemouth bass (Savino and Stein, 1982). Aquatic vegetation is a habitat characteristic associated with Esocidae (Pflieger, 1997), and is widely recognized for its importance to various life stages of northern pike (Inskip, 1982; Bry, 1996). Although the identification of these habitat types is similar to other esocids (Raney, 1942; Crossman, 1962; Ming, 1968), some variation among species does exist. For example, chain pickerel prefer deeper cooler waters during midsummer, but move into shallow weedy areas in the fall when the water cools (Armbruster, 1959). In addition, habitat use may overlap for esocids of similar size (Foster, 1980). Northern pike, in most respects, are a larger counterpart of the chain and grass pickerels, preferring slow moving water with dense vegetation (Pflieger, 1997). More specifically, Casselman and Lewis (1996) reported that larger northern pike were caught at low vegetation densities, and the smallest fish were taken in the densest vegetative cover. Because grass pickerel rarely grow to a size greater than 350 mm, we expected them to have vegetation requirements similar to small northern pike. Muskellunge, like other pikes, also select habitat with clear water and dense growths of aquatic vegetation (Pflieger, 1997).
Our findings in Indiana suggest grass pickerel may occur in communities of varying quality as measured by the Index of Biotic Integrity and the Qualitative Habitat Evaluation Index (QHEI). The Index of Biotic Integrity has been shown in several studies (Sullivan et al., 2004; Lau et al., 2006) to be significantly related to QHEI scores, including this study. However, grass pickerel abundances were not related to IBI scores. In Ohio, grass pickerel have been shown to decrease in numbers or become extirpated in streams where channelizadon has destroyed habitat (White et al, 1975). However, this didn't apply in our study.
Grass pickerel are lie-in-wait predators (Moyle and Cech, Jr., 2004) that are not adapted to fast moving water. Weinman and Lauer (2007) confirmed they are carnivores, while Ming (1968) showed them attacking prey from a place of concealment. Thus, our findings showing a negative relationship of grass pickerel catch with riffle areas was expected, and validates known behavior with habitat use. All grass pickerel in our study were found in low flow areas, generally pools or run areas, and none were found in riffle areas.
Temperature, pH, turbidity, conductivity and dissolved oxygen can have significant impacts on the fish assemblage of lotic systems (Eriksen et al, 1996; Hauer and Hill, 1996; Margeanu et al, 1998). For example, northern pike growth is negatively affected by warm water (Margeanu et al, 1998; Headrick and Carline, 1993), although they can tolerate low oxygen levels (Mecozzi, 1989). The range of water quality factors measured in Indiana showed grass pickerel were able to adapt and survive in a diversity of warm water habitats, including sites with very high temperatures (>32 C) and low dissolved oxygen (
Habitat use of grass pickerel in Indiana was based on two main components: in-stream cover and slow moving water, which are similar for other esocids (Pflieger, 1997) with one exception. Logs/woody debris are not in-stream cover components that have been previously described as habitat used by other esocids, although Rust et al (2002) indicated the importance of wood in Wisconsin lakes to muskellunge spawning and age-0 muskellunge. Grass pickerel may be unique in this aspect as they are the tertiary predator (e.g., primarily consuming fish and crayfish; Weinman and Lauer, 2007), and in small streams with little aquatic vegetation must use other types of cover to stalk prey species. Identifying this grass pickerel habitat need can be used to manage stream ecosystems, particularly when aquatic vegetation is sparse or non existent. Without these structures, the stream trophic structure may be biologically altered from the top down. In addition, because grass pickerel are eury- tolerant of temperature, oxygen and some stream physical habitat components, this fish can survive where other top-predators may not.
Acknowledgments. - We thank the Indiana Department of Environmental Management, the Muncie Bureau of Water Quality and the City of Elkhart for aiding in data collection and Ball State University for funding. We also thank Lorena Edenfield, Joe Foy, Stephen Warmer, Stacey Sobat, Heath Headley, Nate Thomas, Trudy Perkins, Mark Pyron and Adam Cain for assistance in the field and laboratory.
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SUBMITTED 16 JULY 2007 ACCEPTED 1 JANUARY 2008
MICHELLE L. CAIN, THOMAS E. LAUER1 and JAMIE K. LAU
Aquatic Biology and Fisheries Center, Ball State University, Munde, Indiana 47306
1 Corresponding author: Telephone: (765)285 8825; FAX: (765)285 8804; e-mail: firstname.lastname@example.org
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