Analysis of genetic diversity in common loon Gavia immer using RAPD and mitochondrial RFLP techniques

We used random amplified polymorphic DNA (RAPD) and restriction fragment length polymorphisms (RFLP) of mitochondrial cytochrome b (cyt b ) gene to evaluate the genetic diversity in common loon Gavia immer populations from two regions in the United States: New England (NE) and Michigan (MI). RAPD analysis with 18 primers showed 74% polymorphism in NE and 50% in MI loons (similarity coefficient F = 0.92). Although no population-specific markers were found, the frequencies of some RAPD bands varied between the two populations suggesting geographical differences. RFLP analyses with Bam HI enzyme and a 307-bp mitochondrial cyt b gene showed four haplotypes in the NE loon samples and two in the MI samples. The mtDNA haplotype diversity was 0.74 for NE and 0.51 for MI loons, supporting the RAPD data that NE loons have greater genetic diversity than MI loons.     

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     

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.     

A novel method for captive rearing and translocation of juvenile common loons

Common loons (Gavia immer) are diving waterbirds that are particularly challenging to keep in captivity due to their specific behavioral and physiologic needs, special housing requirements, and susceptibility to stress-related disease. We report a novel method for housing and captive rearing common loon chicks that was developed as part of the first-ever loon translocation effort in southeast Massachusetts, from 2015 to 2017. Thirteen loon chicks were reared in aquatic pens in a natural lake environment, utilizing noninvasive feeding and monitoring techniques that avoided human habituation. Chicks were reared in aquatic pens for 16–28 days before being released onto the lake. All chicks remained clinically normal and were monitored on the lake for up to 4 months following release. At least four of the chicks were subsequently confirmed to have survived to adulthood when they returned to the area in breeding plumage two to 3 years following release. Two of these confirmed adults displayed prolonged territorial pair behavior together, and this is an encouraging early sign that captive-reared individuals may form successful breeding pairs in the future. Because most immature loons remain on the ocean until at least 3 years of age, we expect additional captive-reared loons to return to the release area in subsequent years. These husbandry techniques could be applied to other loon and diving bird species that are notoriously difficult to house in captivity. The novel feeding techniques described here could also be adapted for loon chicks being reared in pools or other traditional captive settings.     

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.     

Common Loon Survival Rates and Mercury in New England and Wisconsin

Abstract: Bioaccumulation of toxic environmental mercury may affect the vital rates of piscivores such as the common loon (Gavia immer). Although immediate effects of mercury on early development or reproduction can be determined from short-term field studies or dosing experiments, long-term effects on survival for a long-lived species such as the common loon must be discerned from large, long-term observational data sets. We analyzed band-resight and mercury data for 776 adult loons in Wisconsin and New England, USA, from 1991 to 2001 to 1) estimate annual survival rates and 2) investigate the relation between mercury exposure and survival. The model-averaged estimate of apparent survival was 0.87, whereas the approximate survival rate (accounting for movement) was 0.92. We found no differences in apparent survival by geographic location or sex and no relation between survival and mercury. Power analyses showed that we were only likely to detect differences in survival ≥3%. Small differences in survival (<3%), which may be important to loon population viability, were unlikely to be detected in our dataset. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):665–673; 2008)     

Necropsy findings and environmental contaminants in common loons from New York

Diagnostic and analytical findings are presented for 105 common loons (Gavia immer) found dead or debilitated in New York (USA) from 1972-99. Aspergillosis (23% of cases) and ingestion of lead fishing weights (21%) were the most common pathologies encountered. Stranding on land, shooting, other trauma, gill nets, air sacculitis and peritonitis, and emaciation of uncertain etiology accounted for most of the remaining causes of disease or death. Analysis for total mercury in the liver of 83 loons yielded a geometric mean (gm) of 10.3 mg/kg (wet basis) and range of 0.07 to 371 mg/kg, with emaciated birds generally showing higher levels. Organochlorine contaminant levels in brain were generally low, principally consisting of PCB's (gm = 2.02 mg/kg) and DDE (0.47 mg/kg).     

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.     

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.     

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.     

Nestsite selection by male loons leads to sex-biased site familiarity

1. The concept that animals benefit from gaining familiarity with physical spaces is widespread among ecologists and constitutes a theoretical pillar in studies of territory defence, philopatry and habitat selection. Yet proximate causes and fitness benefits of site familiarity are poorly known. 2. We used data from marked common loons Gavia immer breeding on 98 territories over 14 years to investigate the 'win-stay, lose-switch rule' for nestsite placement (if eggs hatch, reuse nestsite; if predator takes eggs, move nestsite). Males controlled nest placement in this species: pairs used the rule if both members remained the same from the previous nesting attempt or if only the male remained the same but not if only the female remained the same. 3. By means of the nesting rule, male common loons benefited from site familiarity, increasing nesting success by 41% between their first and third years on a territory. In contrast, females exhibited no increase in nesting success with increased territorial tenure. 4. Owing to site familiarity, a male loon competing for a breeding territory faces a considerable 'familiarity deficit' compared with the male breeder already established there. The familiarity deficit probably explains why resident animals often fight hard to retain familiar territories, when challenged, and why animals of many species tend to remain on familiar territories rather than moving when territories of higher intrinsic quality become available nearby.     

Population Growth and Demography of Common Loons in the Northern United States

ABSTRACT We used recent developments in theoretical population ecology to construct basic models of common loon (Gavia immer) demography and population dynamics. We parameterized these models using existing survival estimates and data from long-term monitoring of loon productivity and abundance. Our models include deterministic, 2-stage, density-independent matrix models, yielding population growth-rate estimates (λ) of 0.99 and 1.01 for intensively studied populations in our Wisconsin, USA, and New Hampshire, USA, study areas, respectively. Perturbation analysis of these models indicated that estimated growth rate is extremely sensitive to adult survival, as expected for this long-lived species. Also, we examined 20 years of count data for the 2 areas and evaluated support for a set of count-based models of population growth. We detected no temporal trend in Wisconsin, which would be consistent with fluctuation around an average equilibrium state but could also result from data limitations. For New Hampshire, the model set included varying formulations of density dependence and partitioning of stochasticity that were enabled by the annual sampling resolution. The best model for New Hampshire included density regulation of population growth and, along with the demographic analyses for both areas, provided insight into the possible importance of breeding habitat availability and the abundance of nonbreeding adults. Based on these results, we recommend that conservation organizations include nonbreeder abundance in common loon monitoring efforts and that additional emphasis be placed on identifying and managing human influences on adult loon survival.     

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%).     

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.     

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.     

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.     

Habitat selection by loons in southcentral Alaska

Three species of loons nest in the Matanuska-Susitna Valley, an area 80 km north of Anchorage in southcentral Alaska. This is a region of intense change; its human population doubled between 1980–1990 to almost 36000 people. The Alaska Department of Fish and Game through its Loon Watch Program has monitored 150 to 200 lakes since 1984, half of which are used by loons. Common loons (Gavia immer) nested on lakes of at least 12 ha while Pacific loons (G. pacifica) used lakes as small as 4 ha. Red-throated loons (G. stellata) were found on even smaller ponds. On all waters used by reproductive loons, 89% were connected to other water bodies, 62% of the shore was appropriate for nesting and 85% of the area was <15 ft (4.6 m) deep. This littoral zone comprised habitat for three-spined stickleback fish (Gasterosteus aculeatus), the dominant food source for loons, and supported large macrophyte beds that were used by loons as chick-rearing areas. The effect of human presence on summer loon distribution was dependent on the willingness of humans to accommodate to the needs of loons.