Effect of threadfin shad density on production of threadfin shad and gizzard shad larvae in a Tennessee reservoir

Production of larvae by threadfin shad, Dorosoma petenense, and gizzard shad, D. cepedianum, varied over two orders of magnitude and was regulated by adult threadfin shad abundance over five years in Normandy Reservoir, Tennessee. Significantly more larvae of both species were produced in years following winterkills of threadfin shad (repeated-measures ANOVA, df=4, 75; F > 21.44, p=0.0001). Peak geometric mean catch of threadfin shad larvae in neuston samples was inversely related to biomass (kg ha^–1; r = – 0.91; p=0.031) and density (no. ha^–1; r=– 0.89; p = 0.043) of adult (> 70 mm total length) threadfin shad in mid-summer cove samples. Peak geometric mean catch of gizzard shad larvae was also inversely related to adult threadfin shad biomass (r = – 0.93; p=0.022) and density (r=– 0.88; p=0.046) in cove samples. Winterkills of threadfin shad were size selective, killing all fish under 60 mm total length but allowing some unknown percentage of larger fish to survive. When threadfin shad stocks were reduced by winterkills, surviving threadfin shad and gizzard shad may have taken advantage of less competition for food resources in early spring and increased condition enough to spawn successfully.     

Sediment processing by gizzard shad, Dorosoma cepedianum (Lesueur), in Acton Lake, Ohio, U.S.A.

Gizzard shad are primarily detritivorous in Acton Lake, a 253-ha impoundment in southwestern Ohio, U.S.A. To determine the magnitude of sediment utilization by the gizzard shad population in Acton Lake. I used data on population density and age structure, daily ration, and feeding selectivity in estimating the mass of sediments processed by shad daily from April through November. At densities of 4595–10 645 fish ha⁻¹(wet weight biomass = 90–121 kgha ¹), gizzard shad could process 3.8–23.0 kg of dry sediments ha⁻¹ day ¹. On average throughout the growing season, gizzard shad could process a dry mass of sediments each day equivalent to 13% of shad wet weight biomass. Because of the high rate of sedimentation (> 700 kg dry sediment ha⁻¹ day⁻¹) in Acton Lake, gizzard shad can process < 4% of the freshly deposited sediments each day, and therefore are likely to have little effect on benthic community dynamics in the system.     

Detritivory and the stoichiometry of nutrient cycling by a dominant fish species in lakes of varying productivity

Little is known about the stoichiometry of nutrient cycling by detritivores. Therefore, we explored stoichiometric relationships in an omnivorous/detritivorous fish (gizzard shad, Dorosoma cepedianum) in three lakes that differed in productivity. Gizzard shad can feed on plankton and sediment detritus, but in all three lakes adult gizzard shad derived >98% of carbon (C) and phosphorus (P), and >90% of nitrogen (N) from sediment detritus, and the remainder from zooplankton.
Gizzard shad selectively consumed detritus with higher C, N and P concentrations than ambient lake sediments. Selective detritivory (i.e. the nutrient content of consumed detritus divided by the nutrient content of ambient detritus) was most pronounced in the lake with the lowest detrital nutrient concentrations. N and P cycling rates per fish were also consistently higher in this lake, in agreement with the prediction of stoichiometry theory that excretion rates should increase with food nutrient content. Among-lake differences in nutrient cycling rates were unrelated to inter-lake variation in fish body nutrient contents, which was minimal. The N:P ratio excreted was near Redfield (∼14:1) in all three lakes.
Stoichiometric analyses showed that the C:N and C:P ratios of sediment detritus were much higher (∼2.8×) than ratios of gizzard shad bodies, revealing substantial N and P imbalances between consumers and their food source. Gizzard shad alleviate N imbalance by selectively feeding on high N detritus (low C:N, high N:P), and apparently alleviate P imbalance by excreting nutrients at a higher N:P than that of their food or their bodies. Thus, this detritivore apparently regulates nutrient acquisition and allocation via both pre-absorption processes (selective feeding) and post-absorptive processes (differential N and P excretion).     

Influence of gizzard shad on phytoplankton size and primary productivity in mesocosms and earthen ponds in the southeastern U.S.

Gizzard shad (Dorosoma cepedianum), a filter feeding omnivore, can consume phytoplankton, zooplankton and detritus and is a common prey fish in U.S. water bodies. Because of their feeding habits and abundance, shad have the potential to affect primary productivity (and hence water quality) directly through phytoplankton grazing and indirectly through zooplankton grazing and nutrient recycling. To test the ability of shad to influence primary productivity, we conducted a 16-day enclosure study (in 2.36-m³ mesocosms) and a 3-year whole-pond manipulation in 2–5 ha earthen ponds. In the mesocosm experiment, shad reduced zooplankton density and indirectly enhanced chlorophyll a concentration, primary productivity, and photosynthetic efficiency (assimilation number). While shad did not affect total phytoplankton density in the mesocosms, the density of large phytoplankton was directly reduced with shad. Results from the pond study were not consistent as predicted. There were few changes in the zooplankton and phytoplankton communities in ponds with versus ponds without gizzard shad. One apparent difference from systems in which previous work had been conducted was the presence of high densities of a potential competitor (i.e., larval bluegill) in our ponds. We suggest that the presence of these extremely high larval bluegill densities (20–350 larval bluegill m^–3; 3–700 times higher density than that of larval gizzard shad) led to the lack of differences between ponds with versus ponds without gizzard shad. That is, the influence of gizzard shad on zooplankton or phytoplankton was less than the influence of abundant bluegill larvae. Differences in systems across regions must be incorporated into our understanding of factors affecting trophic interactions in aquatic systems if we are to be able to manage these systems for both water quality and fisheries.     

Lake trophic state and the limnological effects of omnivorous fish

Ecologists have hypothesized that planktivorous fish have greater effects on the plankton and water quality of oligotrophic lakes than eutrophic lakes. We tested this hypothesis in a tank-mesocosm experiment of factorial design in which five biomass levels of filter-feeding omnivorous gizzard shad (Dorosoma cepedianum) were cross-classified with two levels of lake trophic state achieved by filling tank-mesocosms with water and plankton transported by truck from two lakes with different trophic states. The presence of gizzard shad significantly increased total phosphorus, primary productivity, chlorophyll, and particulate phosphorus (PP) 2–20 and 20–200 μm and significantly decreased Secchi depth, cladocerans, copepods and PP > 200 μm. The effects of gizzard shad on chlorophyll, Secchi depth, cladocerans, copepods and PP 2–20 and > 200 μm were dependent on lake trophic state and most intense in the eutrophic lake system. This experiment suggests that filter-feeding omnivorous fish interact synergistically with trophic state so that the limnological effects of omnivorous fish become more intense with increased eutrophication.     

Particulate and filter feeding in threadfin shad, Dorosoma petenense, at different light intensities

Previous workers inferred from stomach analyses that threadfin shad ( Dorosoma petenense ) ate plankton by both filter and particulate feeding. These inferences were confirmed in this study by laboratory experiments in which both types of feeding were observed. Threadfin shad consumed relatively small food particles (≤ 0.39 mm) by filtration, while larger prey (7.5 mm) were eaten individually. The shad were able to filter feed on small foods (brine shrimp nauplii and phytoplankton) at all light intensities from 0 to 9 × 10¹ fL. These data indicate that under natural conditions shad can filter feed at any time of the 24 h period if food conditions are sufficient to trigger feeding. Filter feeding is probably induced by chemoreception rather than vision. The particulate feeding rate decreased as light intensity decreased, reaching a minimum between 9 × 10⁻⁴ and 9 × 10⁻⁵ fL. From these data it is inferred that particulate feeding is a visual process in this species, requiring intensities equivalent to bright moonlight or greater. Filter and particulate feeding abilities allow threadfin shad to consume most of the different types of plankton and to change their diet with seasonal changes in the composition of the plankton.     

Effects of gizzard shad on benthic communities in reservoirs

Effects of gizzard shad Dorosoma cepedianum on benthic communities in a large southern reservoir (Lake Texoma, U.S.A.) were examined during two field enclosure and exclosure experiments in which enclosures were stocked at high and low densities in 1998 and 1999, respectively. In both years, chironomid abundance significantly increased in treatments that excluded large fishes from foraging on sediments. Mean abundance of chironomids and ostracods were significantly higher ( P  < 0·05) in exclosures than enclosures stocked with gizzard shad at 1140–1210 kg ha⁻¹. In 1999, benthic invertebrate abundances did not differ ( P  > 0·08) between exclosure and enclosures stocked at 175–213 kg ha⁻¹. Per cent organic matter, algal abundance and abundance of other macroinvertebrates in sediments did not differ significantly among treatments in either year. Although chironomid abundance was reduced in gizzard shad enclosures in 1998, food habits from this and other studies showed that adult gizzard shad in Lake Texoma only consumed detritus and algae. It is likely that high sedimentation rates in Lake Texoma limit the ability of gizzard shad to regulate algae and detritus in benthic sediments. Thus, it is concluded that disturbance of benthic sediments by gizzard shad caused the observed reduction in chironomid abundance, rather than through consumption or competition for resources.     

Seasonal and interannual variation in nutrient fluxes from tributary inputs, consumer recycling and algal growth in a eutrophic river impoundment

We measured tributary inputs, algal nutrient demand and excretion rates of consumers (gizzard shad and zooplankton) at a eutrophic river impoundment. During two summers with contrasting flow regimes, tributary inputs accounted for 38% (1998) and 3% (1999) of algal N demand and 95% (1998) and 17% (1999) of algal P demand. Gizzard shad contributions averaged 14% and 20% of algal demand for N whereas P contributions were 31% and 58% (1998, 1999; respectively). Zooplankton recycling accounted for a comparable fraction of algal P demand (47%) but a larger fraction of N demand (43%) because their excretia were N rich (N:P = 13:1) compared to fish (7:1). Nutrient release by one of the consumers (gizzard shad) was compared with tributary loading over a nine-year period to assess inter-annual variation in their relative importance. Historical records of inflow chemistry, discharge and gizzard shad biomass showed that variation in tributary inputs was the primary determinant of seasonal and inter-annual variation in nutrient loading. Consumer-derived nutrients were important in late-summer and during years when tributary inputs were low. We propose a conceptual model in which primary production is regulated by external nutrient loading and consumer recycling acts to stabilize and sustain production during periods of diminished external inputs.     

Development of hypoosmoregulatory ability in allis shad Alosa alosa

When different ages of juvenile allis shad Alosa alosa (18–74 days post-fertilization) were exposed for 48 h to elevated salinity (25 or 30), all ages could tolerate moderates salinity (25) whereas only the older stages could tolerate the higher salinity. In 88 days post-fertilization allis shad, acclimatation to full-strength sea water for 2 weeks did not lead to significant changes in total water content and gill Na⁺/K⁺-ATPase activity whereas total Na⁺ content increased within 5 days to values which remained stable thereafter. Together, these data suggest that juvenile allis shad develop a limited euryhalinity, which allows them to fully adapt to hyperosmotic environment immediately after transfer.     

Nutritional importance of detritivory in the growth and condition of gizzard shad in an Ohio reservoir

Synopsis Gizzard shad,Dorosoma cepedianum, in Acton Lake, Ohio, ingested foods of varying nutritional quality during the 1981–1983 growing seasons. Adult (ages 2–4) fish fed on a mixed diet (ORG>30%; C:N<7:1) of zooplankton and organic detritus in early summer, and on detrital materials (ORG<16%; C:N>11:1) during the remainder of the growing season. Age 0 (<35 mm standard length) fish ingested only detrital materials. The nutritional quality (ORG = 10 – 20%; C:N<11:1) of these foods displayed little seasonal variation, but was higher than that of organic detritus taken by adult fish in late summer and autumn. Growth and condition of gizzard shad were poor when the diet consisted of detrital materials; however, age 4 fish (1983) grew rapidly and condition improved when zooplankton were consumed. These results suggest that ingestion of poor-quality detritus can reduce the growth and condition of gizzard shad in Acton Lake, whereas the seasonal inclusion of high-quality zooplankton in the diet can result in rapid growth and improved condition.     

Stable isotope evidence of ontogenetic changes in the diet of gizzard shad Dorosoma cepedianum

Stable sulphur isotopic composition (δ³⁴S) of gizzard shad Dorosoma cepedianum was used to investigate the seasonal and ontogenetic variation in the diet of young and adult fish. This study evaluated fish from a hypereutrophic lake that had recently undergone a 40% reduction of large (>300 mm total length, L _T) D. cepedianum biomass as part of a biomanipulation experiment, which aimed at reducing internal nutrient loading. Dorosoma cepedianum δ³⁴S values showed evidence of ontogenetic changes with young fish (<200 mm L _T) depending more on benthic food sources than adults (>200 mm L _T). The δ³⁴S composition of the adult fish suggested an increasing importance of zooplankton in the diet, although benthic food sources remained part of the diet of all D. cepedianum collected in this study. The results indicated that benthic feeding is used by D. cepedianum of all sizes, suggesting that biomanipulation efforts may need to target all sizes of fish to realize benefits.     

Evaluating short-term effects of omnivorous fish removal on water quality and zooplankton at a subtropical lake

We evaluated a biomanipulation program to test for short-term changes in water quality (chlorophyll a, Secchi depth, total phosphorus) and macrozooplankton biomass following partial removal of omnivorous gizzard shad Dorosoma cepedianum. The removal occurred at a eutrophic subtropical lake, and responses were compared to an unmanipulated control lake using a before-after-control-impact paired series analysis. The removal reduced the biomass of large (>300 mm) gizzard shad by 75% over 2 years via a subsidized commercial gill net fishery. However, the total population biomass of gizzard shad was reduced by approximately 32% from an average pre-manipulation biomass of 224 kg ha⁻¹ due to the size selectivity of the gear, which did not effectively capture small fish (<300 mm). No significant short-term changes in chlorophyll a concentration, Secchi depth, total phosphorus concentration or macrozooplankton biomass were detected following biomanipulation. The partial removal may have fallen short of the biomass reduction required to cause ecosystem responses. Our results suggest that moderate omnivore removals (i.e., <40% biomass reduction) will have little short-term benefits to these lakes, and future manipulations should use a less size-selective gear to achieve a larger total biomass reduction.     

Bird abundance and species richness on Florida lakes: influence of trophic status,lake morphology,and aquatic macrophytes

Data from 46 Florida lakes were used to examine relationships between bird abundance (numbers and biomass) and species richness, and lake trophic status, lake morphology and aquatic macrophyte abundance. Average annual bird numbers ranged from 7 to 800 birds km^–2 and bird biomass ranged from 1 to 465 kg km^–2. Total species richness ranged from 1 to 30 species per lake. Annual average bird numbers and biomass were positively correlated to lake trophic status as assessed by total phosphorus (r = 0.61), total nitrogen (r = 0.60) and chlorophyll a (r = 0.56) concentrations. Species richness was positively correlated to lake area (r = 0.86) and trophic status (r = 0.64 for total phosphorus concentrations). The percentage of the total annual phosphorus load contributed to 14 Florida lakes by bird populations was low averaging 2.4%. Bird populations using Florida lakes, therefore, do not significantly impact the trophic status of the lakes under natural situations, but lake trophic status is a major factor influencing bird abundance and species richness on lakes. Bird abundance and species richness were not significantly correlated to other lake morphology or aquatic macrophyte parameters after the effects of lake area and trophic status were accounted for using stepwise multiple regression. The lack of significant relations between annual average bird abundance and species richness and macrophyte abundance seems to be related to changes in bird species composition. Bird abundance and species richness remain relatively stable as macrophyte abundance increases, but birds that use open-water habitats (e.g., double-crested cormorant, Phalacrocorax auritus) are replaced by species that use macrophyte communities (e.g., ring-necked duck, Aythya collaris).     

Zooplankton–phytoplankton relationships in shallow subtropical versus temperate lakes Apopka (Florida,USA) and Trasimeno (Umbria,Italy)

This study compares and contrasts the dynamics of phytoplankton, zooplankton, and nutrients in two of the largest shallow lakes in the USA (Lake Apopka, Florida) and Europe (Lago Trasimeno, Umbria, Italy) and considers particularly the biomass ratio of zooplankton to phytoplankton (BZ:BP) in relation to nutrient levels and in the context of data from other subtropical and temperate lakes. Lake Apopka is hypereutrophic with higher concentrations of total phosphorus (TP), nitrogen (TN), and nearly an order of magnitude higher BP than Lago Trasimeno. However, combined data from the two lakes can be fit to a single log–log regression model that explains 72% of the variability in BP based on TP. In contrast, BZ has a significant positive log–log relationship with TP only for Lago Trasimeno, and is much lower than expected based on the TP concentrations observed in Lake Apopka. Lake Apopka has a fish assemblage that includes high densities of gizzard shad (Dorosoma cepedianum) and threadfin shad (D. petenense), similar to other eutrophic Florida lakes that also have extreme low BZ. The ratio BZ:BP is below 0.01 in Lake Apopka, 10-fold lower than in Trasimeno and among the lowest values reported in the literature. Although stress of high water temperature and a greater proportion of inedible cyanobacteria may be contributing factors, the collective results support an emerging view that fish predation limits the biomass of crustacean zooplankton in subtropical lakes. Handling editor: S. I. Dodson     

Threadfin shad, Dorosoma petenense Günther, mortality: causes and ecological implications in a South-eastern United States reservoir

Cold stress was identified as an important factor influencing both reservoir-wide mortality and impingement of threadfin shad, Dorosoma petenense , during the period October 1976 to April 1977 in Watts Bar Reservoir, Tennessee. Relative numbers and size frequency of impinged threadfin shad were similar to the relative numbers and size frequency of shad preyed upon by sauger, Stizostedion canadense , and skipjack herring, Alosa chrysochloris . This relationship implies that the factor mainly responsible for impingement, low temperature, also influences prey vulnerability. Threadfin shad made up 99% of the combined diet of sauger and skipjack herring from November until the threadfin shad disappeared in January. These predators did not readily switch to alternative prey in the short term, but by the next autumn 25–100% of the diet was alternative prey. Reappearance of threadfin shad the year following mass mortality and ability of the predators to vary their diet emphasize the resilient nature of some predator-prey systems.     

Effects of environmental factors on survival, growth, and production of American shad larvae

Episodic increases in temperature of 5°C above 20° C, over 48 h or declines in pH of 1·0 unit from pH 7·0 reduced survival of yolk-sac and feeding-stage larvae of American shad Alosa sapidissima . Over 16 days all measures of survival, growth, and production were more favourable at each higher temperature in the 15–25° C range. More favourable responses were also obtained at the higher prey level (500 v . 50 Artemia nauplii l^-1) and at the higher pH (7·5 v . 6·5). Combinations of high temperature and high prey levels, at pH 7·5, led to highest larval production. Little growth or production occurred at 15° C, regardless of pH or prey level. The effect of pH was strong with respect to survival, but weak with respect to growth. In attempts to restore American shad populations by larval stocking, release times and sites can be critical to optimize survival and eventual returns. Releases of larvae potentially will be most effective when made at temperatures >20° C, pH>7·0, and prey levels >50 1^-1. These conditions are most likely to occur in Maryland tributaries of Chesapeake Bay between mid-May and early June.     

Piscivore enhancement effects on food webs depend on planktivore body size and species composition in replicated whole lake experiments

Recent syntheses of trophic cascade and biomanipulation research have suggested that the effects of piscivores on planktivorous fish populations are reduced, when planktivores are capable of outgrowing predator gape limitation and in systems with complex food web interactions. These hypotheses, however, have not been tested in long-term, whole-lake, experiments where processes such as fish recruitment and compensatory food web responses may be important. We conducted a replicated whole-lake experiment to test for the effects of supplemental piscivore introductions on food webs of eutrophic lakes dominated by deep-bodied planktivores. Responses to piscivore enhancement were compared between lakes differing in food web structure due to the presence of omnivorous gizzard shad (Dorosoma cepedianum). A significant decrease in the relative abundance of juvenile planktivorous fish, and an increase in total benthic macroinvertebrate density was observed in lakes containing mainly bluegills (Lepomus machrochirus). In contrast, lakes containing gizzard shad exhibited no significant responses to piscivore manipulation. Our results support the hypothesis that food webs in lakes dominated by deep-bodied planktivorous fish species respond weakly to piscivore enhancement. In addition, our findings support the hypothesis that cascading trophic interactions are weaker in lake ecosystems with more complex food web interactions such as those containing gizzard shad.     

Movement into floodplain habitats by gizzard shad (<Emphasis Type="Italic">Dorosoma cepedianum</Emphasis>) revealed by dietary and stable isotope analyses

Floodplain habitats have been inferred to provide a variety of functions for aquatic organisms, yet few studies have documented movement between channel and aquatic floodplain habitats. We exploited spatial variation in stable isotope ratios of gizzard shad (Dorosoma cepedianum) to document movement between floodplain lakes and the main river channel of the Brazos River, Texas, during a period of frequent hydrologic connectivity. Additionally, we examined stomach contents of shad to determine if ontogenetic diet shifts or faunal exchange best explained variation in isotopic ratios. Regression analysis indicated significant relationships between gizzard shad size and isotopic ratios in oxbow lakes, whereas these relationships were not significant for the main channel. Plots of individual fish in each habitat suggested that adult shad migrated into oxbow lakes during floods whereas juveniles assimilated material produced in oxbows. Some adults in oxbows had signatures similar to juveniles, and these individuals were probably long-term oxbow residents. The proportion of adults with a “river” signature was greater in the oxbow with the shortest flood recurrence interval where opportunities for faunal exchange were more frequent. Analysis of stomach contents indicated almost total overlap between adult and juvenile diets indicating that movement between habitats having different isotopic ratios of basal resources rather than ontogenetic dietary shifts best explained patterns of isotopic variation in Brazos River gizzard shad.     

Stable isotopes of carbon and nitrogen as indicators of diet and trophic structure of the fish community in a shallow hypereutrophic lake

Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes were employed to elucidate energy flows and trophic interactions in Lake Apopka, a hypereutrophic lake in central Florida, U.S.A. Isotope compositions of lake biota ranged from −27·1 to −3·0‰ for δ¹³C, and from 3·7 to 13·9‰ for δ¹⁵N. The food web was based primarily on plankton production with diatoms, Microcystis and zooplankton dominating the diet of fish. Carbon isotope evidence showed that pico- and nano-phytoplankton were not a direct carbon source for fish, but were important to zooplankton. δ¹⁵N mass balance estimates indicated that planktivorous fish obtained 48–85% of their diets from zooplankton. The ∼3‰ range of δ¹⁵N in gizzard shad reflected increasing dependence on zooplankton as fish grew whereas the positive relationship between total length and δ¹⁵N of largemouth bass reflected increasing predation on larger planktivorous fish with growth. The broad ranges of δ¹³C (−25·9 to −9·5‰) and δ¹⁵N (5·8 to 14·4‰) of blue tilapia were indicators of diet diversity. Two presumed omnivores (brown bullhead and white catfish) and piscivores (black crappie, largemouth bass and Florida gar) were found to depend on planktivorous fish. However, stable isotope data revealed no trophic links between blue tilapia, an abundant fish in the near-shore area, and piscivores.     

Population structure and food habits of white crappie Pomoxis annularis Rafinesque in a turbid Oklahoma reservoir

We assessed population structure and food habits of white crappie Pomoxis annuluris Rafinesque in Lake Carl Blackwell, a turbid reservoir in north-central Oklahoma, U.S.A. White crappie [ n = 8549; total length ( t.l. ) mean = 147 mm, s.d. = 28 mm, range = 73–445 mm] were collected with gillnets, frame nets, and hoop nets. Proportional stock density was 2, as 97% of the fish were under 200 mm t.l . Relative stock density was 96, 2, 1, and 1 for 'stock-quality', 'quality-preferred', 'preferred-memorable', and 'memorable-trophy' lengths. Total annual mortality rate was 54% for fish older than 2 years. Growth was poor and lower than the Oklahoma-Arkansas regional averages ( P <0.01). Relative weights were 79, 86, 106, 109, and 123 for fish in the stock-quality, quality-preferred, preferred-memorable, memorable-trophy, and trophy size categories. Ephemeroptera nymphs were an important dietary component for fish 150 mm t.l. or smaller. Gizzard shad Dorosoma cepedianum (Lesueur) was a major forage item, especially for fish greater than 150 mm t.l. ( P <0.01). Empirical information on prey availability indicates that the white crappie population is forage limited and that gizzard shad are not available in appropriate sizes or sufficient numbers to sustain the predator population. Poor fish condition, slow growth, and high mortality are probably the result of inadequate forage.