Ora E. Johannsson

LARGE NONLETHAL EFFECTS OF AN INVASIVE INVERTEBRATE PREDATOR ON ZOOPLANKTON POPULATION GROWTH RATE

We conducted a study to determine the contribution of lethal and nonlethal effects to a predators net effect on a preys 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.

Tribute to R. G. Boutilier: the effect of size on the physiological and behavioural responses of oscar, Astronotus ocellatus, to hypoxia.

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 ṀO2 was 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.

Life History and Productivity of Mysis Relicta in Lake Ontario

Mysis relicta is a common food of many fish species in Lake Ontario, yet little was known about its life history or production. We determined growth rates, cohort development patterns, reproductive patterns, and production of the population at a deep water station over a 4-year period. Four replicate vertical net hauls were collected monthly from April until October, each year. All animals were counted and 100 from each replicate were measured. The original mysid data of N.H.F. Watson, collected in 1971–72, were reanalyzed and the results compared with the present study. Most reproduction occurred in late autumn; the eggs hatched late the following winter; and the offspring reached maturity approximately 18 month later. In 1971 and 1972, abundances were similar to that in 1984 but cohort development was less synchronous. Average growth rate (1984–1987) in the first summer was 0.035 mm.d−1, through the winter it fell to 0.012 mm.d−1, and during the second summer it was 0.029 mm.d−1. Production was 1.90g.m−2 in 1984 and 2.62 g.m−2 in 1986. Examination of sources of error suggest these values might underestimate production by 11% to 27%. Immigration prevented the estimation of production in 1985 and 1987

Roles of Predation, Food, and Temperature in Structuring the Epilimnetic Zooplankton Populations in Lake Ontario, 1981–1986

Abstract We sampled phytoplankton, zooplankton, and alewives Alosa pseudoharengus and measured water temperature in Lake Ontario during 1981–1986. Through the use of general linear regression models we then sought evidence of control of the eplimnetic zooplankton community (mid-July to mid-October) by producers, consumers, and temperature. Our measures of the zooplankton community were total biomass, cladoceran biomass, and the ratio of large to small Daphnia spp. (D. galeata mendotae and D. retrocurva). Zooplankton population variables assessed were abundance, egg ratio, and productivity. Through factor analysis, factors were created from the standardized, transformed independent variables for use in the regression analyses. Regression models showed significant inverse relationships (P < 0.05) between alewives and Bosmina longirostris (abundance, production, and egg ratio), Ceriodaphnia lacustris (egg ratio), and Daphnia retrocurva (egg ratio). Bosmina longirostris and D. retrocurva egg ratios were inver...

Filtering Impacts of an Introduced Bivalve (Dreissena polymorpha) in a Shallow Lake: Application of a Hydrodynamic Model

ABSTRACT Nonindigenous species may exert strong effects on ecosystem structure and function. The zebra mussel (Dreissena polymorpha) has been attributed with profound changes in invaded ecosystems across eastern North America. We explored vertical profiles of water flow velocity and chlorophyll a concentration in western Lake Erie, over rocky substrates encrusted with Dreissena, to assess the extent to which mussels influence coupling between benthic and pelagic regions of the lake. Flow velocity was always low at surveyed sites (less than or equal to 2.9 cm s-1) and declined in direct proximity to the lakebed. Mean chlorophyll a concentration was also low (less than 5μg L-1) at all sites and depths. Chlorophyll a concentration was positively correlated with distance above lakebed and was lowest (0.3μg L-1) directly adjacent to the lakebed. Spatial patterns of zooplankton grazers could not explain observed vertical gradients in chlorophyll concentration. Hydrodynamic modeling revealed that filtering effects of Dreissena in a nonstratified, shallow basin depend mainly on upstream chlorophyll concentration, intensity of turbulent diffusion, feeding efficiency of the mussel colony, and the distance downstream from the leading edge of the mussel colony. In contrast to widespread perceptions that molluscs reduce phytoplankton concentration only adjacent to the lakebed, modeling scenarios indicated that depletion occurs throughout the water column. Depletion was, however, inversely proportional to distance above the lakebed. Simulation results are consistent with field-based observations made in shallow water habitats populated by large Dreissena populations in the Great Lakes and elsewhere. Results from this study indicate that zebra mussels strongly enhance coupling between pelagic and benthic regions in shallow lakes. Enhanced coupling between these regions explains, in part, high population densities of Dreissena and of many benthic invertebrates in ecosystems invaded by zebra mussels.

Spatial and long-term temporal assessment of Lake Ontario water clarity, nutrients, chlorophyll a, and zooplankton

Secchi depth, total phosphorus, soluble reactive phosphorus, silica, chlorophyll a, and zooplankton (density, biomass, and average size) were sampled as part of a lake-wide, seasonal (spring, summer, and fall) assessment of Lake Ontario in 2003 to characterize the status of the lower aquatic food web. For each parameter, spatial comparisons were performed to test for differences between habitats (nearshore and offshore) and between regions (east and west) during each season. Significant differences between habitats were found only for silica and chlorophyll a; silica was higher nearshore in fall, and chlorophyll a was higher offshore in fall. Significant differences between regions were detected in Secchi depth, epilimnetic zooplankton mean length, total phosphorus and Cercopagis pengoi density and biomass; Secchi depth and zooplankton mean length were higher in the east in spring, total phosphorus and Cercopagis pengoi biomass were higher in the west in summer, and Cercopagis pengoi biomass was higher in the east in fall. Cercopagis pengoi was present lake-wide in summer and fall, but Bythotrephes longimanus was present only in fall in the Kingston basin. Mean spring total phosphorus, soluble reactive phosphorus, chlorophyll a, and epilimnetic zooplankton density and biomass are at or near record low levels. As we move into the future, persistent low levels of these lower food web elements will continue to stress alewife populations both through reduced food resources and food quality for zooplankton, and may force these fish to seek alternative food such as Mysis.

The effects of silver on intestinal ion and acid-base regulation in the marine teleost fish, Parophrys vetulus

Exposure to elevated silver (as AgNO3) concentrations (6-9 microM) in seawater was associated with comparably high silver concentrations in the intestinal fluids of the lemon sole (Parophrys vetulus), and a tendency for reduced drinking rate. The effects of silver on intestinal ion and acid-base regulation were studied using in situ perfusion of the intestine. Intestinal net Cl- uptake was reduced from 0.4 to 0.1 and intestinal net Na+ uptake from 0.2 to 0 mmol kg(-1) x h(-1) during silver exposure (9 microM). At the same time, intestinal HCO3- net efflux was reduced from 0.2 to 0.1 mmol kg(-1) x h(-1). Both intestinal Na+ and Cl- uptake and Cl-/HCO3- exchange are thus sensitive to silver, but to different extents. None of the observed effects were reversible during 24 h of recovery. Intestinal water transport was highly variable in vivo in the perfused preparation, and no significant effect of silver exposure was observed. However, in vitro intestine preparations exhibited reduction of intestinal net water flux from 4 to 1 microl cm(-2) x h(-1) during silver exposure together with reduced unidirectional Cl- influx. Reduced water intake and transepithelial water transport in silver-exposed fish resulted in moderate hemoconcentration evident from higher hematocrit values, but not in increased plasma ion levels. The latter could reflect a compensatory response via increased branchial Na+/K+-ATPase levels, observed in silver-exposed fish, indicative of increased branchial ion transport capacity. Impairment of intestinal ion and water transport as a result of silver intake via drinking could be an important part of the fatal cascade of physiological effects observed in marine fish during acute silver exposure.

Age and Growth of Alewives in the Changing Pelagia of Lake Ontario, 1978–1992

Abstract We documented the age and growth of alewives Alosa pseudoharenqus in Lake Ontario during 1978–1992 and determined if growth was affected by intraspecific competition for epilimnetic zooplankton, lake temperature, or demand of salmonine piscivores for prey. Ages of juvenile alewives were determined from scales during 1978–1983, and ages of juvenile and adult alewives were determined from otoliths during 1984–1992. Indices of abundance for alewives were calculated from spring bottom trawl catches in 1978–1992; zooplankton density and epilimnetic temperature were monitored at two stations during 1981–1991; and salmonine demand each year during 1978–1992 was calculated with a simulation model. Although we encountered 11-year-old alewives, few fish lived longer than 7 years, and most fish in the population were younger than 6 years. Mean sizes at ages 1, 2, and 3 in spring averaged 93 mm (5.1g), 133 mm (17 g), and 149 mm (22 g), but from age 3 to age 8, mean size increased by only 5–7 mm and 2–3 g per...

Comparison of Lake Ontario Zooplankton Communities Between 1967 and 1985: Before and After Implementation of Salmonid Stocking and Phosphorus Control

Abstract Two strong, contrasting management strategies were applied to Lake Ontario during the 1970s. There has been a 40 percent reduction in phosphorus loadings to the lake and an exponential increase in salmonid stocking. A comparison of zooplankton community structure and abundance from 1981 to 1985 with that of earlier studies (1967 to 1972) found no detectable change in the range of abundances or community composition between the two periods. Clearly, neither management strategy has had a discernible impact on the zooplankton community to date; however, the potential for change in the system remains high.

Effect of Long-term Fasting on the Use of Fatty Acids as Trophic Markers in the Opossum Shrimp Mysis relicta—A Laboratory Study

ABSTRACT Fatty acids are potential trophic markers to trace feeding relationships in aquatic ecosystems primarily because lipid reserves of organisms broadly reflect dietary sources of lipids and can therefore provide information on the availability of key fatty acids in the food web. However, the use of fatty acids for such applications may be constrained by the degree to which the fatty acid composition of organisms is obscured by factors other than straightforward uptake from the diet. Thus, we studied the effect of long-term fasting, under controlled laboratory conditions, on the lipid content and fatty acid composition of field-caught Mysis relicta. Periods of 3 to 6 weeks are required to induce clear effects of fasting in M. relicta. Relative proportions of docosahexaenoic acid (DHA; 22:6n-3) and arachidonic acid (ARA; 20:4n-6) increased with decreasing lipid contents during fasting. DHA proportions in the total fatty acid composition of fasting M. relicta were significantly higher in comparison to field-caught animals. M. relicta with high proportions of DHA (> 25%) and with low lipid contents (< 14% of dry weight) can be clearly identified as fasting animals. Such thresholds will help to improve the validity of fatty acids in trophic studies.