Nutrient cycling by fish supports relatively more primary production as lake productivity increases.

Animals can be important in nutrient cycling in particular ecosystems, but few studies have examined how this importance varies along environmental gradients. In this study we quantified the nutrient cycling role of an abundant detritivorous fish species, the gizzard shad (Dorosoma cepedianum), in reservoir ecosystems along a gradient of ecosystem productivity. Gizzard shad feed mostly on sediment detritus and excrete sediment-derived nutrients into the water column, thereby mediating a cross-habitat translocation of nutrients to phytoplankton. We quantified nitrogen and phosphorus cycling (excretion) rates of gizzard shad, as well as nutrient demand by phytoplankton, in seven lakes over a four-year period (16 lake-years). The lakes span a gradient of watershed land use (the relative amounts of land used for agriculture vs. forest) and productivity. As the watersheds of these lakes became increasingly dominated by agricultural land, primary production rates, lake trophic state indicators (total phosphorus and chlorophyll concentrations), and nutrient flux through gizzard shad populations all increased. Nutrient cycling by gizzard shad supported a substantial proportion of primary production in these ecosystems, and this proportion increased as watershed agriculture (and ecosystem productivity) increased. In the four productive lakes with agricultural watersheds (>78% agricultural land), gizzard shad supported on average 51% of phytoplankton primary production (range 27-67%). In contrast, in the three relatively unproductive lakes in forested or mixed-land-use watersheds (>47% forest, <52% agricultural land), gizzard shad supported 18% of primary production (range 14-23%). Thus, along a gradient of forested to agricultural landscapes, both watershed nutrient inputs and nutrient translocation by gizzard shad increase, but our data indicate that the importance of nutrient translocation by gizzard shad increases more rapidly. Our results therefore support the hypothesis that watersheds and gizzard shad jointly regulate primary production in reservoir ecosystems.

An analysis of the mechanisms governing species replacements in crayfish.

We investigated mechanisms governing replacement of the native crayfish Orconectes sanborni by an invading cryafish, Orconectes rusticus. The two species had similar life histories, habitat preferences, and feeding patterns in allopatric and sympatric stream areas. Orconectes rusticus young-of-year (YOY) grew faster than O. sanborni YOY in the field. Adult O. rusticus were larger and, hence, dominant over adult O. sanborni; YOY were non-aggressive. In laboratory experiments, adult crayfish (about 28 mm carapace length or larger) were not susceptable to predation by largemouth bass (Micropterus salmoides, 30 cm total length) and did not alter shelter use when fish were present. Orconectes rusticus YOY were less susceptible to predation than O. sanborni YOY. Orconectes rusticus YOY reduced their vulnerability to largemouth bass by occupying shelters more often than YOY O. sanborni. In mixed-species mateselection experiments, male O. rusticus and male O. sanborni preferentially mated with O. rusticus females. Inappropriate mate selection in sympatry may have caused the 90% reduction in recruitment for both species in 1982. Orconectes rusticus probably maintains greater population growth than O. sanborni, because (1) more gravid O. rusticus females occurred in sympatry, (2) O. rusticus produced more young than O. sanborni, and (3) O. rusticus young grew faster. Reproductive interference, acting synergistically with differences in aggressive dominance and young-of-year susceptibility to predation, appears to serve as the major mechanisms regulating replacement of O. sanborni by O. rusticus in Ohio streams.

Complex predator-prey interactions and predator intimidation among crayfish, piscivorous fish, and small benthic fish.

Predator-prey interactions were studied among a small prey fish (the johnny darter Etheostoma nigrum) and two predators (crayfish Orconectes rusticus and smallmouth bass Micropterus dolomieui) with complementary foraging behaviors. When only smallmouth bass were present, darters reduced activity to 6% of control rates and spent most of the time hiding under tile shelters. When only crayfish were present, darter activity and shelter-use were similar to controls. When both crayfish and bass were present, an interaction occurred. Darters, normally inactive in the presence of bass, were often forced to move by approaching crayfish and thus activity increased to 19% of control rates. Also, darters were often evicted from shelters by intruding crayfish. Thus, crayfish increased the vulnerability of small fish to bass by evicting them from shelters and causing increased activity. Conversely, bass increased the vulnerability of small fish to crayfish by forcing these fish to seek cover under shelters occupied by crayfish. Intimidation effects of bass on darters last for some time. After a 30-min exposure to bass, darters showed reduced activity and increased shelter use lasting at least 24 h after the bass was removed. Thus predators, throught intimidation, can influence prey behavior even though the predators are no longer present.

Bluegill growth as modified by plant density: an exploration of underlying mechanisms.

Bluegill (Lepomis macrochira) growth varies inconsistently with plant density. In laboratory and field experiments, we explored mechanisms underlying bluegill growth as a function of plant and invertebrate density. In the laboratory, bluegills captured more chironomids (Chironomus riparius) than damselflies (Enallagma spp. and Ischnura spp.), but energy intake per time spent searching did not differ between damselfly and chironomid treatments. From laboratory data, we described prey encounter rates as functions of plant and invertebrate density. In Clark Lake, Ohio, we created 0.05-ha mesocosms of inshore vegetation to generate macrophyte densities of 125, 270, and 385 stems/m2 of Potamogeton and Ceratophyllum and added 46-mm bluegill (1/m2). In these mesocosms, invertebrate density increased as a function of macrophyte density. Combining this function with encounter rate functions derived from laboratory data, we predicted that bluegill growth should peak at a high macrophyte density, greater than 1000 stems/m2, even though growth should change only slightly beyond 100 stems/m2. Consistent with our predictions, bluegills did not grow differentially, nor did their use of different prey taxa differ, across macrophyte densities in the field. Bluegills preferred chironomid pupae, which were relatively few in numbers but vulnerable to predation, whereas more cryptic, chironomid larvae, which were associated with vegetation but were relatively abundant, were eaten as encountered. Bluegills avoided physid snails, which were abundant. Contrary to previous work, vegetation did not influence growth or diet of bluegill beyond relatively low densities owing to the interaction between capture probabilities and macroinvertebrate densities.