We conducted a study to determine the contribution of lethal and nonlethal effects to a predator's net effect on a prey's population growth rate in a natural setting. We focused on the effects of an invasive invertebrate predator, Bythotrephes longimanus, on zooplankton prey populations in Lakes Michigan and Erie. Field data taken at multiple dates and locations in both systems indicated that the prey species Daphnia mendotae, Daphnia retrocurva, and Bosmina longirostris inhabited deeper portions of the water column as Bythotrephes biomass increased, possibly as an avoidance response to predation. This induced migration reduces predation risk but also can reduce birth rate due to exposure to cooler temperatures. We estimated the nonlethal (i.e., resulting from reduced birth rate) and lethal (i.e., consumptive) effects of Bythotrephes on D. mendotae and Bosmina longirostris. These estimates used diel field survey data of the vertical gradient of zooplankton prey density, Bythotrephes density, light intensity, and temperature with growth and predation rate models derived from laboratory studies. Results indicate that nonlethal effects played a substantial role in the net effect of Bythotrephes on several prey population growth rates in the field, with nonlethal effects on the same order of magnitude as or greater (up to 10-fold) than lethal effects. Our results further indicate that invasive species can have strong nonlethal, behaviorally based effects, despite short evolutionary coexistence with prey species.
We examined stressors that have led to profound ecological changes in the Lake Ontario ecosystem and its fish community since 1970. The most notable changes have been reductions in phosphorus loading, invasion by Dreissena spp., fisheries management through stocking of exotic salmonids and control of sea lamprey (Petromyzon marinus), and fish harvest by anglers and double-crested cormorants (Phalacrocorax auritus). The response to these stressors has led to (i) declines in both algal photosynthesis and epilimnetic zooplankton production, (ii) decreases in alewife (Alosa pseudoharengus) abundance, (iii) declines in native Diporeia and lake whitefish (Coregonus clupeaformis), (iv) behavioral shifts in alewife spatial distribution benefitting native lake trout (Salvelinus namaycush), threespine stickleback (Gasterosteus aculeatus), and emerald shiner (Notropis atherinoides) populations, (v) dramatic increases in water clarity, (vi) predation impacts by cormorants on select fish species, and (vii) lake trout recruitment bottlenecks associated with alewife-induced thiamine deficiency. We expect stressor responses associated with anthropogenic forces like exotic species invasions and global climate warming to continue to impact the Lake Ontario ecosystem in the future and recommend continuous long-term ecological studies to enhance scientific understanding and management of this important resource. 490Résumé : On trouvera ici un examen des facteurs de stress qui ont modifié profondément l'écosystème du lac Ontario et sa communauté de poissons depuis 1970. Les changements les plus importants ont été la réduction de l'apport de phosphore, l'invasion des Dreissena spp., la gestion de la pêche, notamment l'empoissonnement de salmonidés exotiques et le contrôle de la grande lamproie marine (Petromyzon marinus), ainsi que la récolte des poissons par les pêcheurs sportifs et les cormorans à aigrette (Phalacrocorax auritus). La réaction à ces facteurs a eu pour conséquen-ces: (i) le déclin de la photosynthèse des algues et de la production du zooplancton épilimnétique, (ii) la diminution de l'abondance du gaspareau (Alosa pseudoharengus), (iii) la réduction des Diporeia indigènes et des grands corégonesCan.
We compared predatory demand by pelagic planktivorous prey fish with invertebrate production in Lake Michigan during 1987 and in Lake Ontario during 1990. Predation by the planktivores in Lake Ontario was nearly fourfold higher than in Lake Michigan Qapprox. 87 g wet ~e i g h t -r n -~-~e a r -' ) .Predation rates on Mysis were comparable in Lakes Michigan and Ontario (approx. 21 g-m-2-year-'), while predation on Diporeia was markedly higher in Lake Michigan than in Lake Ontario (21.3 vs. 8.5 g wet ~e i g h t . m -~.~e a r -' ) .In Lake Ontario, predatory demand on zooplankton exceeded our best estimate of production by a factor of 1.7. Similarly, predation estimates on Mysis in Lake Ontario were 1.2-2.0 times the estimated rate of production, depending on the production model used. Lake Michigan planktivores consumed approximately 55% of available zooplankton production in 1987, indicating that competition for prey resources, if operating, was not as intense as that in Lake Ontario in 1990. It is unclear how to resolve the paradox that predation could markedly exceed available prey production in some cases. There could be sources of error in the estimates of both the supply and demand sides of these trophic relationships.RCsnmC : Nous avons compare B la production d'invertCbrCs la demande predatrice des poissons pelagiques planctivores servant eux-rnemes de proies, en 1987 dans le lac Michigan et en 1990 dans le lac Ontario. Dans ce dernier lac, la prkdation par les planctivores Ctait prks de quatre fois plus ClevCe que dans le lac Michigan (environ 87 g poids frai~-rn-~.an-'). Les taux de predation 5 17&gard des Mysis 6taient comparables dans les deux lacs (environ 21 g.ms2.an-'1, tandis que la prddation i lqCgard des Diporeia etait nettement plus haute dans le lac Michigan que dans le lac Ontario (21,3 contre $ 3 g poids frai~-m-~-an-l). Dans le lac Ontario, la demande des prkdateurs i l'kgard du zooplancton dkpassait d'un facteur de 1,7 notre meilleure estimation de la production.Be meme, les estimations de la prCdation a l'kgard des Mysis dans le lac Ontario correspondait B 1,2-2,0 fois le taux estimC de production, selon le modkle de production retenu. Les planctivores du lac Michigan ont consommC environ 55% de la production disponible de zooplancton en 1987, ce qui indique que la concurrence 8 l'Cgard des ressources em proies, si elle existait, n'itait pas
Mysis relicta and planktivorous fish in Lakes Ontario and Michigan both feed on crustacean zooplankton in the metalimnion. Are these zooplankton sufficient to meet the energy requirements of mysids? Could mysids remove a significant proportion of zooplankton production? Are the energy requirements in the two lakes similar? Comparisons in Lake Ontario of zooplankton consumption based on clearance rates, with energy requirements based on bioenergetic modelling, revealed that individual mysids, particularly larger ones, required additional energy sources. The denser mysid populations beyond 100 m depth in Lake Ontario could exert high mortality rates on metalimnetic zooplankton, remove a significant proportion of daily zooplankton production, and consequently, potentially compete with planktivorous fish. At depths < 100 m, the mysid population was smaller and could remove only 6–19% of zooplankton production per day in summer when competition with planktivorous fish would be maximal. Generation time is shorter in Lake Michigan because winter growth rates remain high. Consequently, less energy is required to complete a generation in Lake Michigan than in Lake Ontario.
Stable isotope analysis of the potential prey and predator can be combined with gut content analysis to quantify the diet. This dietary knowledge allows the quantitative assessment of the role of key species in energy and contaminant transfer, their impact on prey communities, and their susceptibility to perturbation. The diet of Mysis relicta was examined in Lake Ontario in spring, summer, and autumn using both techniques. Mysids fed on the bottom during the day and in the pelagia and on the bottom at night. A trophic fractionation of 2.2‰ N between mysids and their prey provided the best correspondence between the observed stable isotope signature of mysids and that estimated from their diet. Tissue turnover rate of d 13 C was slow compared with that of d 15 N. Diatoms formed 50% of the assimilated diet in May. In September, 25% of large mysids feeding on the bottom contained amphipod parts and 20% contained phytoplankton. The remainder of the diet consisted of zooplankton and rotifers. The contribution of amphipods and phytoplankton could not be quantified. Revised daily consumption estimates, based on this new diet information and clearance rate estimates of consumption, gave daily consumption estimates similar to those estimated from previous bioenergetic modelling.
SUMMARY The physiological and behavioural responses of two size groups of oscar(Astronotus ocellatus) to hypoxia were studied. The physiological responses were tested by measuring ṀO2 during decreasing environmental oxygen tensions. Larger oscars were better able to maintain oxygen consumption during a decrease in PO2, regulating routine ṀO2 to a significantly lower PO2 threshold (50 mmHg)than smaller oscars (70 mmHg). Previous studies have also demonstrated a longer survival time of large oscars exposed to extreme hypoxia, coupled with a greater anaerobic enzymatic capability. Large oscars began aquatic surface respiration (ASR) at the oxygen tension at which the first significant decrease in ṀO2was seen (50 mmHg). Interestingly, smaller oscars postponed ASR to around 22 mmHg, well beyond the PO2 at which they switched from oxyregulation to oxyconformation. Additionally, when given the choice between an hypoxic environment containing aquatic macrophyte shelter and an open normoxic environment, small fish showed a greater preference for the hypoxic environment. Thus shelter from predators appears particularly important for juveniles, who may accept a greater physiological compromise in exchange for safety. In response to hypoxia without available shelter, larger fish reduced their level of activity (with the exception of aggressive encounters) to aid metabolic suppression whereas smaller oscars increased their activity, with the potential benefit of finding oxygen-rich areas.
The extent to which temperature, temperature gradients, predator smell, and prey availability influence the migratory behaviors and vertical distribution of the opossum shrimp, Mysis relicta, was explored through controlled laboratory experiments and comparisons with field distributions of mysids in Lake Ontario. By varying environmental conditions in 2-m tall experimental columns in a temperature-controlled room, we determined that mysids prefer temperatures between 6uC and 8uC with limited movement into waters of 12uC or higher. No mysids moved into waters above 16uC in the absence of prey. However, a higher proportion of mysids moved into temperatures of 14uC and 16uC (but not 18uC) when densities of Daphnia pulicaria exceeding 120 L 21 were present at those temperatures. Mysis avoided waters with kairomones from a primary mysid predator, the alewife (Alosa pseudoharengus). The rate of temperature change with depth did not restrict mysid movements. A temperature preference function based on the experimental data was applied to an existing model of mysid vertical distribution. The modified model predicted the depth of maximum mysid density to within 1 m and yielded high percentage overlap index values when compared with published mysid vertical distributions in Lake Ontario. Our approach may be used to model how diurnal, seasonal, and larger climactic changes can impact both the vertical position and feeding ecology of mysids, a keystone species in many deep-water pelagic food webs.
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