Factors influencing productivity of common loons (Gavia immer) breeding on circumneutral lakes in Nova Scotia, Canada

Common loons (Gavia immer) are top predators that are sensitive to biotic and abiotic conditions associated with their breeding lakes, so factors such as lake chemistry and human activity or disturbance are thought to influence their seasonal and long-term reproductive success. We used two indices of loon productivity to evaluate (1) temporal patterns and (2) relationships with physical and chemical lake characteristics and human activities. Data collected from 1991 to 2000 by volunteers of the Canadian Lakes Loon Survey (CLLS) in Nova Scotia showed that loon productivity, as indexed by both the proportion of resident pairs that produced at least one large young (P_s1) and the proportion of successful pairs that produced two large young (P_s2), did not vary substantially from year to year and showed no linear trend from 1991 to 2000. Average estimates (1991–2000) for P_s1 and P_s2 were 0.49 ± 0.02 and 0.43 ± 0.03, respectively, and the mean number of chicks per residential pair over that time was 0.75 ± 0.04. We found that human disturbance and shoreline development did not influence loon productivity during the prefledging stage on lakes surveyed by CLLS volunteers. Proportion of resident pairs rearing at least one large young was independent of dissolved organic carbon (DOC) concentrations of breeding lakes, but there was a positive relationship between the proportion of successful pairs rearing two large young and DOC. Both indices of loon productivity tended to be negatively correlated with lake pH. These results were not consistent with other findings that loon productivity generally declines with lake acidity, but likely reflect the preponderance of circumneutral (pH 6.5–7.0) lakes surveyed by the CLLS volunteers in Nova Scotia.     

Bioaccumulation of mercury in yellow perch (Perca flavescens) and common loons (Gavia immer) in relation to lake chemistry in Atlantic Canada

Mercury biomagnifies in aquatic foodwebs in freshwater lakes, and common loons (Gavia immer) breeding in eastern Canada can be exposed to reproductively toxic concentrations of mercury in their fish prey. We assessed the bioaccumulation and biomagnification of mercury in juvenile and adult common loons, and their preferred prey: yellow perch (Perca flavescens) in Kejimkujik National Park (KNP), Nova Scotia by measuring mercury levels and stable isotope ratios in tissues. Total mercury levels and stable-carbon (δ¹³C) and nitrogen isotope ratios (δ¹⁵N) were determined in composite whole-fish samples from lakes in KNP and blood samples from juvenile and adult loons captured on lakes in KNP and southern New Brunswick. Geometric mean mercury concentrations were 0.15 and 0.38 μg/g (wet wt.) in small (9-cm fork length) and large (17-cm fork length) yellow perch, and were 0.43 and 2.7 μg/g (wet wt.) in blood of juvenile and adult common loons, respectively. Mercury concentrations in perch and loons were positively associated with body mass and δ¹⁵N values. Juvenile loons and large yellow perch had similar mercury levels and δ¹⁵N values, indicating similar trophic status despite their 22-fold difference in body mass. Mercury concentrations were higher in yellow perch and common loons in acidic lakes. Our findings highlight the importance of both chemical and ecological factors in understanding mercury biomagnification in lakes and associated risks to fish-eating wildlife. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

The oral bioavailability and toxicokinetics of methylmercury in common loon (Gavia immer) chicks

We compared the toxicokinetics of methylmercury in captive common loon chicks during two time intervals to assess the impact of feather growth on the kinetics of mercury. We also determined the oral bioavailability of methylmercury during these trials to test for age-related changes. The blood concentration-time curves for individuals dosed during feather development (initiated 35 days post hatch) were best described by a one-compartment toxicokinetic model with an elimination half-life of 3 days. The data for birds dosed following completion of feather growth (84 days post hatch) were best fitted by a two-compartment elimination model that includes an initial rapid distribution phase with a half-life of 0.9 days, followed by a slow elimination phase with a half-life of 116 days. We determined the oral bioavailability of methylmercury during the first dosing interval by comparing the ratios of the area under the blood concentration-time curves (AUC(0--> infinity )) for orally and intravenously dosed chicks. The oral bioavailability of methylmercury during the first dosing period was 0.83. We also determined bioavailability during both dosing periods using a second measure because of irregularities with intravenous results in the second period. This second bioavailability measure estimated the percentage of the dose that was deposited in the blood volume (f), and the results show that there was no difference in bioavailability among dosing periods. The results of this study highlight the importance of feather growth on the toxicokinetics of methylmercury.     

An outbreak of type E botulism among common loons (Gavia immer) in Michigan's upper peninsula

An epizootic of type E botulism (Clostridium botulinum) occurred among common loons (Gavia immer) along the Lake Michigan shore of Michigan's Upper Peninsula (USA) during October and November 1983. An estimated 592 dead loons washed ashore along the Garden Peninsula. Type E botulinal toxin was demonstrated in blood samples and stomach contents of dead loons, and in samples of three species of dead fish found on the Lake Michigan shore. We suspect that loons acquired botulism by ingesting sick or dead fish containing type E toxin.     

Antimicrobial resistant Salmonella spp. isolated from double-crested cormorants (Phalacrocorax auritus) and common loons (Gavia immer) in Florida.

Antimicrobial resistant Salmonella spp. were found in double-crested cormorants and common loons in Florida. Single or multiple resistance occurred in all Salmonella agona isolates from cormorants, primarily to ampicillin, sulfonamids, streptomycin, neomycin, and kanamycin. Similar patterns of resistance were found in S. agona isolates from common loons. In addition, isolates of S. Saint paul from loons were found resistant to tetracycline and streptomycin, while 2 of 7 isolates of S. infantis were resistant to tetracycline only.     

Responses of breeding common loons to human activity in upper Michigan

Breeding populations of the common loon (Gavia immer) in Michigan have declined in the past several decades, resulting in classification of this species as threatened under state law. Factors responsible for the decline are unknown, but may include toxic contaminants, mortality in commercial fish nets, and human disturbance of breeding sites. To assess the latter possibility, 960 hours of observation were devoted to observing human-loon interactions on two sets of lakes, one with restricted human use (minimal or no shoreline development and open to canoe use only), and another with unrestricted use (varying amounts of shoreline development and motorboat traffic). Six mated pairs of loons on six restricted use (r-u) lakes were compared to eight loon pairs on seven open-use (o-u) lakes. The number of nests that hatched young per nest started was not significantly different between the two sets of lakes (7 of 13 (0.62) on r-u lakes vs 8 of 17 (0.47) on o-u lakes), despite significantly more human activity on o-u lakes. Chicks hatched per pair of loons were likewise not significantly different (1.1 vs 1.2 on r-u and o-u lakes, respectively). Fledging success was significantly lower on r-u lakes (7 chicks fledged of 11 hatched) than on o-u lakes (13 fledged of 13 hatched). Human activity on o-u lakes was 2–3 times that on r-u lakes during chick rearing, but time spent by adult loons tending and feeding chicks was not significantly different between the two types of lakes. The larger size of most o-u lakes may have allowed loons a greater opportunity to avoid human disturbance. Higher levels of human activity did not affect production of chicks by loons under the conditions observed, but these results should not be extrapolated to lakes experiencing much higher human use.     

Activity budgets of a marked common loon (Gavia immer) nesting population

A newly devised nightlighting technique was used to capture breeding adult common loons (Gavia immer) at the Seney National Wildlife Refuge in northern Michigan in 1989. The behaviors of 6 pairs of known-sex, color-marked common loons were subsequently quantified during the breeding cycle in 1990. Collected observational data indicate that foraging, resting, locomotion, and preening were frequent throughout the breeding cycle. Time spent foraging was greatest during the pre-nesting period (53 to 57%), but declined significantly during the nesting and post-nesting periods (p<0.05). Time spent foraging during the pre-nesting period was similar to that of fall and winter studies. During the pre-nesting period adult loons spent about 15% of the time in locomotion; this was significantly greater than the other time periods (p<0.05) and is attributed to selecting a nest site. During the nesting cycle, almost half of each bird's time spent was nest-sitting. Sexual differences were negligible during nest-sitting. Resting and chick-rearing were the predominant behaviors during the post-nesting period and were responsible for the biggest difference in parental duties. Time spent preening declined from 8% during the pre-nesting period to 4 to 5% during the post-nesting period. Time spent by nesting pairs to produce chicks is approximately 10% during pre-nesting, 48 to 49% during nesting, and between 38 to 44% during post-nesting. By quantifying and establishing behavioral standards, subtle abnormalities or changes can be detected to better manage for viable common loon populations.     

Chlorophyll-phosphorus relationship in acidified lakes of the High Tatra Mountains (Slovakia)

Concentrations of total phosphorus (TP) and chlorophyll a (CHLA) were measured in 28 lakes in the High Tatra Mountains (Slovakia) from 1983 to 1990. The relationship between log CHLA and log TP in the Tatra lakes is similar to relationships developed for lakes in other regions, but variation is higher. A part of this variation is caused by acidification of the lakes. In the lakes with pH between 4.9 and 6.3 the CHLA concentrations are often extremely low while TP concentrations decreased, but not as drastically.     

Feeding behavior and modeled energetic intake of common loon (Gavia immer) adults and chicks on small lakes with and without fish

We examined the behavior of common loons, Gavia immer (Brünnich), breeding on small, shallow lakes in central Alberta, Canada that were naturally fishless or contained only small-bodied fishes (minnow lake). For both lake types, adults spent >90% of their time on the nesting lake and >50% of their time foraging. Adult loons on fishless lakes dove more frequently, but dives were of shorter duration than loons on lakes with fish. On two intensively studied fishless lakes, adults fed chicks macroinvertebrates, particularly leeches, whereas on a focal minnow lake, fish made up >70% of prey items delivered by adults. Chicks >36 days of age on a minnow lake spent >50% of their time foraging, whereas older chicks on fishless lakes were highly dependent on food provisioning by adults. Models based on observed foraging patterns indicated that prey size was a better predictor of success in meeting energetic requirements than was feeding behavior (e.g., dive rate, dive success). For most models, estimated energetic intake was higher for loons on minnow lakes than on fishless lakes. Our behavioral observations and model results are consistent with surveys in central Alberta that indicate that breeding Common Loons frequently establish territories on small lakes, but that chicks hatched on lakes completely lacking fish rarely fledge and only if sufficient large invertebrates such as leeches are available.     

Duckling response to changes in the trophic web of acidified lakes

We reared American Black Duck (Anas rubripes Brewster) and Common Goldeneye (Bucephala clangula Linnaeus) ducklings on two Quebec laurentian lakes in which we manipulated brook trout populations (Salvelinus fontinalis Mitchill), lake acidity and lake productivity to relate waterfowl foraging to trophic status of lakes. We developed a preliminary model to assess the effects of lake acidity and productivity, fish predation and interspecific fish/duck competition in relation to available food (aquatic invertebrates). We then validated the model using a factorial analysis of the relationships between the variables pertaining to the diet of the fish and ducklings, and the environmental characteristics of the lakes (acidity, biological production and fish predation).The first factorial axis can be interpreted in terms of biological productivity, while the second axis illustrates the effect that fish have on the quantity and type of food available to ducklings. Two different trends appear to occur depending on whether the carrying capacity of the lake is reduced by acidification of the water or increase through liming or fertilization. In the first case, fish predation appears to have a marked effect on available food, whereas in the second case, interspecific fish/duck competition is apparently to blame for changes in the diet of ducklings. In both instances, but to a lesser extent, fish compete increasingly (exploitation and/or interference) with the ducklings, forcing them to feed to a greater extent in riparian sites that are less accessible to fish.     

Limnological research on northern Apennine lakes (Italy) in relation to eutrophication and acidification risk

Substantial variability was found in the water chemistry of 22 northern Apennine lakes. In a group of lakes there is evidence of disturbance linked to eutrophication processes. Other lakes showed weak ion concentrations and alkalinity below the acidification risk threshold. However no acidified lakes were found. The lack of waterbodies with severely altered hydrochemistry may explain why no clear relationship between plankton community structure and water chemistry was observed.     

Susceptibility to freshwater acidification by two species of loon: Red-throated Loon (Gavia stellata) and Arctic Loon (Gavia arctica) in southwest Sweden

In southwest Sweden, the two species of loon, Gavia stellata and G. arctica, have shown different trends in population size and production of young during the last decades. Both species fish in oligotrophic freshwaters, susceptible to acidification. The number of breeding sites occupied by G. stellata has been reduced by almost 50% during the last 40–50 years. For G. arctica, there are no indications of significant declines in population size or reproductive success during the last 20 years. The different trends in numbers and production of young might reflect different susceptibility to the ecological changes in acidified lakes. G. stellata prefer fishing lakes with high abundance of Acerina cernua and salmonid and cyprinid fish, such as Coregonus albula and Rutilus rutilus. They also feed their prefledged chicks almost entirely on cyprinid and salmonid fish. G. arctica prefer fishing lakes with high transparency and, when feeding in groups, high abundance of Perca fluviatilis. Their young can be fed on aquatic insects as a supplement to the fish diet. Thus, G. stellata to a higher degree than G. arctica relies on fish which are susceptible to low pH, and G. arctica may also benefit from the increased abundance of aquatic insects in lakes with reduced predation from fish. Furthermore, high water transparency is important for the selection of lakes by G. arctica but not by G. stellata. In G. stellata, high contents of mercury in eggs can be related to the intake of fish in lakes affected by acidification.     

Testing hypotheses of social gatherings of common loons (<Emphasis Type="Italic">Gavia immer</Emphasis>)

Three hypotheses are tested to explain the function of common loon social gatherings: Cooperative Foraging, Familiarity, and Reconnaissance. From 1993 to 1999, I studied social gatherings through behavioral observations in Michigan, Wisconsin and Maine. There was no or little evidence for the Cooperative Foraging Hypothesis. Partial or indirect evidence for the Familiarity Hypothesis included the following: (1) Social gatherings lasted both longer and occurred more frequently later in the breeding season (2) Approximately 25% of all the social gatherings observed occurred on neutral territories, and (3) Social gatherings consisted of the same individuals on consecutive days. Predictions from the Reconnaissance Hypothesis were also supported in that a large proportion of individuals participating in social gatherings were non-breeders and that the number of social gatherings observed were not equally distributed across loon territories, but instead increased on territories that had recently undergone a divorce. No one hypothesis was adequate to explain social gatherings and more observations on uniquely marked individuals are needed to further substantiate these initial findings.     

Mercury in tigerfish (Hydrocynus vittatus, Castelnau), green happy (Sargochromis codringtonii, Boulenger) and kapenta (Limnothrissa miodon, Boulenger) from Lake Kariba, Zimbabwe

Concentrations of total mercury were determined in Hydrocynus vittatus (Castelnau), Sargochromis codringtonii (Boulenger), and Limnothrissa miodon (Boulenger) from two localities in Lake Kariba, Zimbabwe.
The mean concentrations of total mercury in H. vittatus from Basin 5 and Basin 2 were 0.08 mg kg⁻¹ and 0.094 mg kg⁻¹, respectively. In S. codringtonii, mean concentrations were 0.004 mg kg⁻¹ and 0.026 mg kg⁻¹ for Basins 5 and 2, respectively. No mercury was detected in L. miodon from Basin 2 while samples from Basin 5 had a mean concentration of 0.069 mg kg⁻¹ (wet weight). Total mercury concentrations were also determined on a dry weight basis.
Within each sampling area, total mercury concentrations were significantly different among species ( P  < 0.05). For H. vittatus and S. codringtonii, total mercury concentrations (in the same species) were not significantly different between the two localities ( P  < 0.05).
The factors causing the observed differences in total mercury between similar species from different localities and among different species in the same locality (sampling area) are discussed. From the observed low levels of mercury in all three species, it was concluded that the mercury constituted 'background levels'. These levels are below the maximum concentrations permissible in human fish foods.     

Population responses of perch (Perca fluviatilis) and roach (Rutilus rutilus) to recovery from acidification in small Finnish lakes

Decreasing trends in atmospheric emissions and acidic deposition during the 1990s have resulted in chemical recovery from acidification in the sensitive surface water systems of southern Finland. Responses of perch and roach populations to the improved water quality were studied in 30 small lakes with the aid of water chemistry monitoring data gathered in 1987–2002 and the data collected from two consecutive periods of gillnet test fishing, 1985–1988 and 2001–2002. In the most acidified lakes, alkalinity and ANC have increased and sulphate and labile aluminium concentrations decreasæed markedly. The response of perch populations to the improved water quality is seen in improved reproduction success, indicated by a higher CPUE in numbers and a lower mean weight. The growth rate of perch has declined as the population density has increased. Roach populations have not recovered in the same way as perch, there being no major changes in NPUE or mean weight. Lower growth rates were, however, observed in the roach populations of all study lakes. The increased perch population density as an obstacle to the recovery of roach populations is discussed. Despite the chemical and biological recovery of the study lakes, the buffer capacity of many headwater lakes is low and the lakes will be sensitive to any increases in acidic deposition in the future.     

Cellular responses of the skin of carp (Cyprinus carpio) exposed to acidified water

The skin of carp was examined after exposure to acidified water. Degenerative cells were common in the upper epidermal layers. During the first days most of these cells exhibited signs of necrosis. Later on the incidence of necrosis decreased and that of apoptosis increased. In the acid-exposed fish, the upper filament cells and pavement cells produced secretory vesicles of high electron density, some of which showed peroxidase activity. This enzyme activity was also present in the glycocalyx covering these cells, and in the cytoplasm of apoptotic cells. Mitotic figures and newly differentiating mucous cells were common in the outer epidermal layers. Mucous cells became elongated and produced mucosomes of high electron density. Mucosomes with peroxidase activity were also found. Club cells increased in number. Chloride cells and solitary chemo-sensory cells, not seen in the controls, appeared in the upper epithelial layer. The skin was invaded by many leucocytes and by pigment-containing cytoplasmic extensions of melanocytes. Some leucocytes apparently penetrated into the club cells. These structural observations reflect the complexity of the physiological response of the skin to acid water.     

The ultrastructure of chloride cells in the gills of the teleostOreochromis mossambicus during exposure to acidified water

Summary Branchial chloride cells, which actively take up ions in the gills of freshwater fish, were studied in tilapia (Oreochromis mossambicus) exposed to sublethally acidified freshwater. Structural damage of cells, resulting in cell death by necrosis, only occurred transiently, when the reduction of water pH was acute rather than gradual. The most prominent effects of water acidification were the rapid increase in the number of chloride cells and the changes in frequency of the different stages of the chloride cell cycle. In the opercular inner epithelium, a twofold increase in cells occurred 48 h after gradual acidification. Cell density stabilized after 4 weeks at a level 5 times that of control fish. Four transitory stages were distinguished in the chloride cell cycle: accessory or replacement cells, immature, mature, and degenerating (apoptotic) cells. In control fish, mature chloride cells dominated (over 50%) with immature and apoptotic cells totalling about 40%. After 4 weeks in acid water, only 13% of the cells were mature. Immature and apoptotic cells dominated, each representing about 40% of the total number of chloride cells. Mature cells apparently age rapidly under these conditions. Thus, chloride cells turn over quickly in acid water, with a minor increase in ion transport capacity of the gills. This conclusion is supported by the observation that opercular and branchial Na⁺/K⁺ ATPase activities in treated fish are only 40%–50% higher than in controls.     

Spatial and temporal changes of benthic macroinvertebrate assemblages in acidified streams in the Bohemian Forest (Czech Republic)

Outflows from two atmospherically acidified lakes in the Bohemian Forest were studied in 2005 and 2007. While Lake ?ertovo has been strongly acidified (～pH 4.6), Lake Laka was only slightly acidified in the past and is recovering now (～pH 5.2). The water chemistry and macrozoobenthos composition were analysed along longitudinal gradients of both lake outflows to determine the present status of their streams. A certain progression in stream chemistry to more neutral conditions was observed along the longitudinal gradients of both streams. However, a possible recovery of macrozoobenthos was evident only in the Lake Laka outflow, mainly via an increasing number of Ephemeroptera and Trichoptera taxa, and an increasing number of Gammarus fossarum, both along the longitudinal gradient and during the period of study. In contrast, no considerable changes were observed in the macrozoobenthos composition of the Lake ?ertovo outflow, presumably because its chemistry was harmful for acidosensitive taxa such as Ephemeroptera and Trichoptera. Plecoptera and Chironomidae were the most numerous groups in this stream. The biological recovery of both streams will depend on further chemical improvement in their catchments as well as on the dispersal ability of benthic organisms.