Prey Selection and Diets of Bluegill Lepomis Macrochirus with Differing Population Characteristics in Two Nebraska Natural Lakes
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Fisheries Management and Ecology, 2003, 10, 31–40 Prey selection and diets of bluegill Lepomis macrochirus with differing population characteristics in two Nebraska natural lakes N.W.OLSON*,C.P.PAUKERT&D.W.WILLIS Department of Wildlife and Fisheries Sciences, South Dakota State University, SD, USA J. A. KLAMMER Nebraska Game and Parks Commission, Bassett, Nebraska, USA Abstract Environmental prey samples and stomach contents of bluegill Lepomis macrochirus (Rafinesque) were collected in spring and summer 2000 from two Nebraska Sandhill lakes. Watts Lake contained a low-density bluegill population, whereas Cozad Lake contained a high-density bluegill population. Bluegill diets from both lakes were compared to determine if bluegill prey preference differed between the two populations. The highest median per cent (by calories) of zooplankton in the diet was 1.3%; the remainder was macroinvertebrates. Watts Lake bluegills preferred (based on Manly’s alpha) amphipods in spring and chironomids in summer. Cozad Lake bluegills did not show a strong preference (compared with Watts Lake) for any macroinvertebrates, but still utilized amphipods and chironomids during both seasons. Larger bluegills in Watts Lake preferred chironomids in summer, but Cozad Lake bluegills did not exhibit this relationship. The higher density Cozad Lake bluegill population appeared to be more opportunistic than the lower density Watts Lake population, but both preferred macroinvertebrates. KEYWORDS: bluegill, diets, food habits, Lepomis macrochirus, Nebraska. 1974; O’Brien, Slade & Vinyard 1976). In addition, Introduction feeding habits may be related to bluegill population Bluegill Lepomis macrochirus (Rafinesque) food habits density (Kitchell, O’Neill, Shutgart, Magnuson & have been well documented throughout its geographical Booth 1974). When prey density is held constant, range, incorporating zooplankton (Mittelbach 1981; higher density bluegill populations typically have Harris, Galinat & Willis 1999), macroinvertebrates slower growth (Weiner & Hanneman 1982), presum- (Schramm & Jirka 1989; Dewey, Richardson & Zigler ably because of intraspecific competition for food 1997), and small fish (Flemer & Woolcott 1966; resources. Higher density fish populations may exhibit Applegate, Mullen & Morais 1967; Engel 1988). Their less selective feeding habits, apparently because they diets may differ temporally (Gerking 1962; Keast & need to forage on less-preferred prey (Schindler, Welsh 1968; Keast 1978). Larger bluegills typically Hodgson & Kitchell 1997). Therefore, we compared select larger prey (Hall, Cooper & Werner 1970); and the food habits and prey selection of two contrasting Engel (1988) found that larger bluegills (i.e. >210 mm) Nebraska Sandhill bluegill populations to determine if primarily preyed upon larger aquatic insects and even differences in bluegill density affected their food habits fish fry. and prey selection. We hypothesized that the higher Optimal foraging theory suggests that bluegill will density Cozad Lake bluegill population may exhibit select for prey items based on the net energy gain, more opportunistic feeding compared with the low- which is affected by prey type, searching time for prey, density Watts Lake bluegill population, which would habitat type, and predator avoidance (Werner & Hall specifically select preferred prey items. Correspondence: Craig Paukert, Southwest Biological Science Center, U.S. Geological Survey, 2255 North Gemini Drive, Flagstaff, AZ 86001, USA (e-mail: [email protected]). *Present address: Montana Cooperative Fishery Research Unit, Department of Ecology, Montana State University, Bozeman, Montana, USA Ó 2003 Blackwell Publishing Ltd 31 32 N. W. OLSON ET AL. Study area Materials and methods Watts Lake and Cozad Lake are both located in the Bluegills were collected along the vegetated, nearshore Sandhills of north-central Nebraska (USA), approxi- zones of Watts Lake in May (i.e. spring) and August mately 80 km apart. Both lakes are relatively shallow (i.e. summer) 2000 and from Cozad Lake in April (i.e. (mean depth: 1.3 m for Watts Lake; 1.9 m for Cozad spring) and August (i.e. summer) 2000. Spring water Lake), are highly vegetated (72% coverage for Watts temperatures in Watts and Cozad Lakes were 17 and Lake and 95% coverage for Cozad Lake), and relat- 13 °C, respectively, whereas summer water tempera- ively clear (Secchi depth readings: 122 cm for Watts tures were 25 °C in both lakes. Daytime electric fishing Lake and 213 cm for Cozad Lake). Although both was used in spring; however, short-term (<4 h) sets of lakes were shallow and vegetated, groundwater springs modified fyke nets (i.e. trap nets) were used in summer prevented winterkills. Both lakes were moderately because of the difficulty in collecting bluegills in these productive, with chlorophyll a values of 2.7 lgL)1 in lakes by electric fishing. All bluegills collected in Cozad Lake and 7.5 lgL)1 in Watts Lake (Paukert & summer were from modified fyke nets. When possible, Willis 2000). Watts Lake is 93 ha in surface area while five bluegills per centimetre length group were collected Cozad Lake has an area of 32 ha. Both catchments are to obtain a diversity of fish lengths. primarily grasslands and have no agricultural crops. Bluegills were weighed (g), total length measured Angling pressure and harvest are presumably minimal, (TL; mm), and stomach contents removed nonlethally based on these lakes being in remote locations and using an acrylic tube (Kamler & Pope 2001) and were poor access (Paukert, Willis & Gabelhouse 2002). subsequently released alive. Stomach contents were Watts Lake contained a low-density bluegill popula- preserved in 10% formalin and returned to the tion compared with Cozad Lake. The mean total laboratory for identification. Laboratory procedures number of bluegills collected per trap (i.e. modified consisted of examining the entire stomach sample for fyke) net night was 32 (n ¼ 10 nets, SE ¼ 4.1) in Watts diet items under a dissection microscope with 40· Lake and 233 (n ¼ 10 nets, SE ¼ 29.0) in Cozad Lake magnification. Prey species were identified and counted (Paukert & Willis 2000). Watts Lake bluegills reach a only if the entire organism or prominent body feature mean length of 168 mm by age 3 whereas Cozad Lake (i.e. heads) was present. Because the use of other body bluegills only attained 126 mm by age 3. Watts Lake features (e.g. spines, segments, carapaces, segmented bluegills also had slightly higher body condition than legs, etc.) would not accurately reveal a quantitative did Cozad Lake bluegills (Paukert & Willis 2000). Both number of a specific organism, they were not included lakes contained largemouth bass, Micropterus sal- in laboratory analyses. All diet items were identified to moides (Lace´ pe` de), black bullhead, Ameiurus melas the class, order, or family level and prey frequency of (R), and yellow perch, Perca flavescens (Mitchill). occurrence was calculated (Bowen 1996). Because Black crappie, Pomoxis nigromaculatus (Lesueur), were caloric content may better estimate energetic content also present in Cozad Lake; Watts Lake contained of diets compared with other methods (Hyslop 1980), saugeye, Stizostedion vitrium (M) · S. canadense the analysis was conducted using per cent of the diet by (Smith). Although they were not sampled in our study, calories. muskellunge, Esox masquinongy (M) were last stocked To estimate environmental prey abundance, in 1997 and are presumably present in Watts Lake in zooplankton and benthos (i.e. macroinvertebrates) low numbers (Nebraska Game and Parks Commission, samples were collected at six locations in each lake unpublished data). However, largemouth bass appear each season. These collections were along the same to be the primary predator on bluegill in Sandhill lakes transect as bluegill sampling immediately following the (Paukert, Willis & Klammer 2002). Both lakes had fish collections. Two composite benthic grabs were moderately abundant populations of largemouth bass, taken at each site with a 156-cm2 Ekman dredge, sieved with the number of 200 mm and longer largemouth in the field, and preserved in 10% formalin. Zooplank- bass collected per hour of electric fishing being 38 ton samples were collected using a 2-m integrated tube (n ¼ 12 stations, SE ¼ 4) in Cozad Lake and 96 sampler, sieved through a 65-lm Wisconsin plankton (n ¼ 12 stations, SE ¼ 13) in Watts Lake (Paukert & net and preserved in 10% formalin. All invertebrate Willis 2000). In addition, there was little evidence that samples were returned to the laboratory, identified to bluegill interspecific competition with other fish species the lowest possible taxon and enumerated. (e.g. yellow perch, black bullhead, and black crappie) For prey items in the stomach and the environment, was detrimental to bluegill growth, condition, or stock weight-length regressions obtained from Dumont, Van structure (Paukert & Willis 2000). de Velde & Dumont (1975) and Smock (1980) were Ó 2003 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2003, 10, 31–40 BLUEGILL FOOD HABITS 33 used to calculate dry weight values. These values were (range, 88–193, r ¼ 0.258, n ¼ 31, P ¼ 0.161). However, then converted to calorie content using equations from relative weight did increase with fish length for the few Cummins & Wuycheck (1971). To estimate bluegill fish collected in August (range, 69–117, r ¼ 0.825, prey selection, Manly’s alpha for constant prey pop- n ¼ 14, P ¼ 0.0003). The weight of individual macro- ulations (Chesson 1978; Krebs 1989) was