Occupancy of yellow‐billed and Pacific loons: evidence for interspecific competition and habitat mediated co‐occurrence

Interspecific competition is an important process structuring ecological communities, however, it is difficult to observe in nature. We used an occupancy modelling approach to evaluate evidence of competition between yellow‐billed (Gavia adamsii) and Pacific (G. pacifica) loons for nesting lakes on the Arctic Coastal Plain of Alaska. With multiple years of data and survey platforms, we estimated dynamic occupancy states (e.g. rates of colonization or extinction from individual lakes) and controlled for detection differences among aircraft platforms and ground survey crews. Results indicated that yellow‐billed loons were strong competitors and negatively influenced the occupancy of Pacific loons by excluding them from potential breeding lakes. Pacific loon occupancy was conditional on the presence of yellow‐billed loons, with Pacific loons having almost a tenfold decrease in occupancy probability when yellow‐billed loons were present and a threefold decrease in colonization probability when yellow‐billed loons were present in the current or previous year. Yellow‐billed and Pacific loons co‐occurred less than expected by chance except on very large lakes or lakes with convoluted shorelines; variables which may decrease the cost of maintaining a territory in the presence of the other species. These results imply the existence of interspecific competition between yellow‐billed and Pacific loons for nesting lakes; however, habitat characteristics which facilitate visual and spatial separation of territories can reduce competitive interactions and promote species co‐occurrence.     

Post-breeding densities,population sizes and lake size partitioning of loon species in western Chukotka,Russia

Loons (family Gaviidae) breed in small ponds and lakes across Arctic landscapes and are high level predators in the lake ecosystems. As such, they may serve as sentinel species, warning humans of alterations in habitat and ecosystem integrity in a region that is undergoing vast change due to climate warming. Here, we characterized the abundance and habitat use of four arctic breeding species of loons in the plains and surrounding mountains of western Chukotka, Russia. Loon surveys were conducted on foot and by boat from 2009–2015. Loon species differed in their use of the four lacustrine habitat types within the study area. In yedoma habitat, the yellow-billed loon (Gavia. adamsii) was the most abundant (0.593 birds/km²); on fluvial plain habitat, Pacific loons (G. pacifica) outnumbered other loons (0.701 birds/km²); mountain valleys were inhabited similarly by pacifica (0.354 birds/km²) and red-throated loons (G.stellata; 0.307); and maritime tundra was used only by pacifica (1.13) and Arctic loons (G. arctica; 0.553). G. adamsii was not observed in mountain valleys or maritime tundra. Mountainous portions of rivers were predominantly occupied by stellata and pacifica, and lowland rivers by stellata, pacifica and arctica. There was a significant difference in the size of lakes occupied by the four congeners. The largest loon, adamsii, occupied the largest lakes (0.69 km²), 80% larger than lakes utilized by pacifica (0.39 km²) and arctica (0.38 km²), and 35 times larger than stellata (0.02 km²). Most lakes were occupied by a single loon species (125/162, 77.2%).     

Comparative nest survival of three sympatric loon species breeding in the Arctic

Identifying factors influencing nest survival among sympatric species is important for understanding and managing sources of variation in population dynamics of individual species. Three species of loons nest sympatrically in northern Alaska and differ in body size, life history characteristics, and population trends. We tested the effects of competition, nest site selection, and water level variations on nest survival of Pacific Gavia pacifica, yellow‐billed G. adamsii, and red‐throated loons G. stellata on the Arctic Coastal Plain in Alaska. Although overall nest survival rates did not differ between species, the factors influencing nest survival varied. Nest site selection influenced nest survival for Pacific and yellow‐billed loons, with both species having high nest survival when nesting on islands and peninsulas, likely due to a reduction in access by terrestrial predators. However, on mainland shorelines, Pacific loons had lower nest survival than yellow‐billed loons, and used a higher proportion of vegetation mats for nest sites suggesting that their smaller body size makes them less adept at nest defense. Nest site selection did not influence nest survival of red‐throated loons corresponding to our result of no nest site preferences by this species. Initiation date had a strong influence on nest survival for Pacific and yellow‐billed loons with nests laid earlier having higher survival. Pacific and yellow‐billed loon nests were susceptible to flooding due to precipitation, which contrasted with red‐throated loons that nest on smaller lakes with lower water level variations. Competition did not affect nest survival for any of the species likely due to most territorial encounters occurring prior to incubation. The only influence we found on red‐throated loon nest survival was differences among years. Our results indicate that loons chose nest sites based on predation risk and that factors influencing breeding success of closely related species may differ under similar breeding conditions.     

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.     

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.     

Potential risk to common loons (Gavia immer) from methylmercury exposure in acidified lakes

Piscivorous birds and mammals in areas remote from point sources of Hg contamination may be exposed to dietary methylmercury concentrations that are sufficiently high to cause reproductive impairment. Common loons (Gavia immer) were observed to show aberrant nesting behavior and low overall reproductive success when Hg concentrations in prey (small fish and crayfish) averaged > 0.3 µg g^–1 wet weight (Barr, 1986), levels known to occur in fish from many lakes in central Ontario. We used data on Hg in Ontario fish to estimate the proportion of lakes where fish small enough for loons to eat (< 250 g) had Hg concentrations that exceeded estimated thresholds for reproductive impairment. Up to 30 % of lakes exceeded thresholds for reproductive impairment, depending on the species of fish and the threshold Hg concentrations chosen. There was a significant negative correlation between fish-Hg concentration and lake pH in most fish species examined. For these species, reductions in sulfate deposition rates are predicted to result in a corresponding reduction of lakes in Ontario having fish with potentially toxic concentrations of Hg.     

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.     

Territory occupancy by common loons in response to disturbance,habitat, and intraspecific relationships

Structure and distribution of animal territories are driven by a variety of environmental and demographic factors. A peninsular population of common loons (Gavia immer) nests on lakes in northwestern Montana, but does not occupy all apparently suitable breeding territories, suggesting unexplained limitations on population growth. To evaluate territorial dynamics of breeding loons in Montana, we created and tested occupancy models that evaluated the hypothesized effects of disturbance, habitat, and intraspecific relationships on territory occupancy by common loons in Montana from 2003 to 2007. Model-averaged results indicated that the abundance of feeding lakes within 10 km (i.e., forage quality) and the number of territorial pairs within 10 km (i.e., density of loons) were equally supported and related to probabilities of occupancy. We found substantial support that the population was in a state of equilibrium, with the numbers of occupied territories stable in time, but not space. We also found that density of territorial pairs was related to the likelihood that an existing territory would be abandoned, but did not influence the establishment of new territories, suggesting the presence of territorial pairs could be a stronger indicator of territory quality to loons than physical lake characteristics. Our index of human disturbance was not well-supported compared to other factors. Our results suggest management for stable or growing loon populations could be achieved using long-term monitoring and protection of occupied territorial lakes and nearby feeding lakes, because these factors most influenced the probability of occupancy of surrounding lakes. © 2011 The Wildlife Society.     

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.     

Reproductive Advantages for Common Loons Using Rafts

Abstract: Artificial nesting islands, or rafts, are often deployed in common loon (Gavia immer) breeding territories to decrease negative impacts of mammalian predation and water-level fluctuations on nesting success. The management value of rafts has been demonstrated in other studies; however, no published studies have quantified the use or associated reproductive benefits of rafts on lakes exhibiting water-level fluctuations. These lakes constitute a major portion of loon nesting habitat in New England and the Midwest. We used long-term data sets from loon survey and raft management efforts on lakes with stable (SWL) and fluctuating water levels (FWL) in New Hampshire and Maine, USA, to compare raft-use patterns on both types of lakes. We then modeled the influence of percentage of nesting attempts on rafts, lake fluctuation type, and human development index on nesting success as a function of the number of nesting attempts. Loons used 76% of all rafts for nesting, and initial use patterns were similar between SWL and FWL lakes. Half (51%) of rafts used for nesting were first used during the initial year of deployment and 90% of those used were used by the third year. Based on our model, we would expect to see an 8.6% increase in nesting success associated with each successive categorical increase in raft use (0–33%, 33–60%, 60–100%). Nesting success varied with lake fluctuation type, increasing by 21.4% from FWL to SWL types. Our model estimated a 12.8% decrease in nesting success associated with an increasing human development index. Naturally nesting loons on FWL lakes are likely to display mean nesting success levels lower than those needed to sustain populations. We suggest that natural nesting habitat on lakes with fluctuating water levels during the loon nesting season may constitute an ecological trap warranting consideration of raft management. Findings in this study are germane for managing breeding loon populations, particularly those on reservoirs requiring permits from the Federal Energy Regulatory Commission.     

Patterns of lake occupancy by fish indicate different adaptations to life in a harsh Arctic environment

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Effects of spatial disturbance on common loon nest site selection and territory success

The common loon (Gavia immer) breeds during the summer on northern lakes and water bodies that are also often desirable areas for aquatic recreation and human habitation. In northern New England, we assessed how the spatial nature of disturbance affects common loon nest site selection and territory success. We found through classification and regression analysis that distance to and density of disturbance factors can be used to classify observed nest site locations versus random points, suggesting that these factors affect loon nest site selection (model 1: Correct classification = 75%, null = 50%, K = 0.507, P < 0.001; model 2: Correct classification = 78%, null = 50%, K = 0.551, P < 0.001). However, in an exploratory analysis, we were unable to show a relation between spatial disturbance variables and breeding success (P = 0.595, R² = 0.436), possibly because breeding success was so low during the breeding seasons of 2007–2008. We suggest that by selecting nest site locations that avoid disturbance factors, loons thereby limit the effect that disturbance will have on their breeding success. Still, disturbance may force loons to use sub-optimal nesting habitat, limiting the available number of territories, and overall productivity. We advise that management efforts focus on limiting disturbance factors to allow breeding pairs access to the best nesting territories, relieving disturbance pressures that may force sub-optimal nest placement. © 2011 The Wildlife Society     

Abundances of waterbird species on lakes in Argentine Patagonia as a function of season,lake size and the presence of mink

Regular counts from 2005 to 2009 were made of the waterbirds inhabiting lakes and ponds in Lanín National Park in the southwestern part of Neuquén Province, Argentina, a landscape dominated by Andean–Patagonian wetlands and forests. Bird surveys conducted on 21 wetland areas detected 8,311 individuals belonging to 27 species from 9 families. The most abundant and frequent species were Ashy-headed Goose (Chloephaga poliocephala), Speckled Teal (Anas flavirostris), Red-gartered Coot (Fulica armillata) and White-winged Coot (Fulica leucoptera). The presence and abundance of bird species with respect to wetland surface area was examined. Small lakes (<100 ha) had higher numbers of individuals, but some species such as the Neotropic Cormorant (Phalacrocorax olivaceus), Great Grebe (Podiceps major), Flying Steamer Duck (Tachyeres patachonicus) and Spectacled Duck (Anas specularis) were more abundant at larger lakes. Speckled Teal, Red Shoveler (Anas platalea) and Chiloe Wigeon (A. sibilatrix) were more common in small- and medium-sized lakes. Most waterbird species were found at a smaller percentage of the lakes where mink were present than at mink-free lakes. Although the ponds and lakes studied are protected within the network of this Argentine National Park, such protection is not implemented effectively. Hence, their future conservation faces several potential threats such as American mink expansion, tourism, fishing and hunting. This information could contribute to the development of management guidelines for the effective conservation of Patagonian wetlands.     

Associations between water chemistry and fish community composition: a comparison between isolated and connected lakes in northern Sweden

1. The correlation between water chemistry, physical variables and fish community composition was examined in 40 small (≤30 ha) coastal lakes in northern Sweden. Twenty of the 40 lakes were isolated from other water bodies and 20 were connected to the Baltic Sea. Lakes were fished in summer, using three different methods. Water chemistry was sampled in late winter prior to ice‐out and pH was measured additionally in summer. 2. Our central question was whether water chemistry plays a greater role in the composition of fish communities in isolated lakes than in connected lakes, as isolated lakes cannot be recolonised once a species has become extinct. 3. Results indicate that winter anoxia affects community composition only in isolated lakes, whereas acidity is of importance in both connected and isolated lakes. Methane (indicating anoxia), was significantly correlated with variation in fish community composition in isolated lakes, and a group of variables that indicate anoxia (CH₄, pCO₂, inorganic carbon and dissolved oxygen) explained 24–34% of the variation. pH alone explained 12% of the variation in community composition for connected lakes and a group of variables indicating acidity (summer and winter pH and ANC) explained 10–20% of the variation in isolated lakes. Lake area was the most important physical variable, being significantly correlated with the variation in fish community composition in connected lakes. 4. In isolated lakes, the presence of pike (Esox lucius), perch (Perca fluviatilis) and roach (Rutilus rutilus) was associated with low CH₄. The occurrence of crucian carp (Carassius carassius) and roach was positively correlated with pH, and the crucian carp was apparently also affected by predation by pike and perch. In connected lakes the effect of anoxia was low, probably due to the possibility of recolonisation and pockets of oxygenated water, allowing pike and perch to persist and thereby limiting the distribution of crucian carp.     

Capturing Common Loons during prenesting and nesting periods

ABSTRACT Several techniques have been used to capture Common Loons (Gavia immer), but effectiveness is limited during periods of the breeding season when loons do not have chicks. From 2005 to 2008, we studied loons in northern Wisconsin and used night lighting to capture loons on nests and also designed a lift net for capturing loons prior to nesting. At night, incubating loons were approached by boat and, when within about 30–60 m, we focused a spotlight on the loon and, once at the nest, captured loons using a landing net. Using this technique, we captured 23 loons in 29 attempts (79%). In addition, taped calls and loon decoys were used to entice prenesting, territorial loons into a shoreline‐based, lift‐net trap at a capture efficiency of 67% (10 captures in 15 attempts) during the second year of use. Our diurnal lift‐net trap and night‐light nest‐capture techniques allowed us to capture adult Common Loons during periods of the breeding season when previous investigators have found loons difficult to catch. These techniques may also be useful for capturing other species of territorial waterbirds, especially other species of loons.     

Using maximum entropy to predict the potential distribution of an invasive freshwater snail

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Fish-eating bird abundance in oligotrophic lakes in Kejimkujik National Park,Nova Scotia,Canada

Aquatic bird population data in 40 oligotrophic lakes and ponds in Nova Scotia, Canada indicates that only lakes > 20 ha support territorial pairs of Common Loon (Gavia immer) and only lakes 40 ha produce their young. Broods of Common Merganser (Mergus merganser americanus) occurred in lakes > 25 ha and loon and merganser broods together occurred only in lakes 80 ha. The fish production in the lakes was estimated from the total phosphorus vs fish yield relationship obtained in similar, oligotrophic lakes in Ontario. Considering the fish consumption and mergansers until fledging along with the maintenance of the adults during the same period (about 200 and 130 kg fish respectively) and the estimated fish production in these lakes, there is a close balance between the size of water body and its fish production to the occupancy and production of piscivorous birds.     

Use of boreal forested wetlands by Pacific loons (Gavia pacifica Lawrence) and horned grebes (Podiceps auritus L.): relations with limnological characteristics

Our objective was to determine if the occurrence and abundance of Pacific loons (Gavia pacifica Lawrence) and horned grebes (Podiceps auritus L.) on 123 wetlands of Yukon Flats National Wildlife Refuge in east central Alaska were related to the limnological characteristics of those wetlands. Aquatic bird-wetland use surveys were conducted in conjunction with limnological sampling from May through September 1985–87 and May through August 1989. Results from logistic regression analysis demonstrated a significant association between the probability of wetland use by Pacific loons and shoreline length, water color, calcium and total phosphorus. Wetland use by horned grebes was related to shoreline length, pH, and chlorophyll. Aquatic bird abundance was then used as a Poisson response variable and modeled as a function of wetland limnological characteristics. Our results indicate that Pacific loon abundance was adequately modeled by linear and quadratic functions of shoreline length, color, pH, calcium and total phosphorus. Horned grebe abundance could not be modeled with this approach. The statistical techniques known collectively as generalized linear models provided a framework for the development of models for aquatic bird use of wetlands. Our results, however, indicate that while this approach shows promise, a better understanding of how to model aquatic bird abundance is needed. We then identify problems in model development and suggest avenues for future research.Contribution of the Missouri Agricultural Experiment Station Journal Series No. 11604.     

Methane sources in arctic thermokarst lake sediments on the North Slope of Alaska

The permafrost on the North Slope of Alaska is densely populated by shallow lakes that result from thermokarst erosion. These lakes release methane (CH₄) derived from a combination of ancient thermogenic pools and contemporary biogenic production. Despite the potential importance of CH₄ as a greenhouse gas, the contribution of biogenic CH₄ production in arctic thermokarst lakes in Alaska is not currently well understood. To further advance our knowledge of CH₄ dynamics in these lakes, we focused our study on (i) the potential for microbial CH₄ production in lake sediments, (ii) the role of sediment geochemistry in controlling biogenic CH₄ production, and (iii) the temperature dependence of this process. Sediment cores were collected from one site in Siqlukaq Lake and two sites in Sukok Lake in late October to early November. Analyses of pore water geochemistry, sedimentary organic matter and lipid biomarkers, stable carbon isotopes, results from CH₄ production experiments, and copy number of a methanogenic pathway‐specific gene (mcrA) indicated the existence of different sources of CH₄ in each of the lakes chosen for the study. Analysis of this integrated data set revealed that there is biological CH₄ production in Siqlukaq at moderate levels, while the very low levels of CH₄ detected in Sukok had a mixed origin, with little to no biological CH₄ production. Furthermore, methanogenic archaea exhibited temperature‐dependent use of in situ substrates for methanogenesis, and the amount of CH₄ produced was directly related to the amount of labile organic matter in the sediments. This study constitutes an important first step in better understanding the actual contribution of biogenic CH₄ from thermokarst lakes on the coastal plain of Alaska to the current CH₄ budgets.