Signals from seabirds indicate changing biology of capelin stocks

We examined the predator-prey interaction between an apex seabird predator, the common murre Uria aalge, and capelin Mallotus villosus, the primary forage fish in the Northwest Atlantic. Sampling of parental deliveries to murre chicks was carried out during the breeding sea- son on Funk Island, located off northeast Newfoundland, Canada. Concurrent vessel surveys were conducted throughout the murre's diving and foraging range around the colony to charac- terize the prey field. Results indicated that in years when capelin was abundant in the size range consumed by murres (suitable capelin), murres delivered large and small fish in similar propor- tions, whereas they delivered more large fish when suitable capelin abundance was low. Consid- ering the relative abundances of small and large suitable capelin, these observations suggest neg- ative prey switching by the predator. Using foraging theory, we derived a model which estimates the probability of delivering a specific prey type (large or small capelin or other prey) to the chick based on prey availabilities. This quantitative model was capable of reproducing the general pat- terns in the observations. It also allowed estimating the shape of the common murre's multispecies functional response (MSFR) which indicated that this would conform to the definition of prey switching, and could then be classified as a Type 3. From an applied perspective, our results support the use of predator diets as indicators of their food base, but also highlight the need for understanding the shape of the predator's MSFR for quantitative development of these types of applications.

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 501:279-290 (2014) - DOI: https://doi.org/10.3354/meps10696 Underwater behaviour of common murres foraging on capelin: influences of prey density and antipredator behaviour Kevin A. Crook, Gail K. Davoren* Department of Biological Sciences, University of Manitoba, Room 212B Biological Sciences Building, 50 Sifton Rd, Winnipeg, Manitoba R3T 2N2, Canada *Corresponding author: gail.davoren@ad.umanitoba.ca ABSTRACT: Diving behaviour of seabirds has been studied using data logging devices, but little is known about underwater predator–prey interactions during dives. We used stationary video cameras to investigate how the underwater foraging behaviour of common murres Uria aalge was influenced by the density and behaviour of their main prey fish, capelin Mallotus villosus, at spawning sites on the northeast Newfoundland coast during July, 2009-2012. From ~720 h of video, we analyzed 99 events where capelin and murres were observed together, ranging from 1-20 s, and 952 events where murres were observed alone, ranging from 1-14 s. Although 91% of all video footage of capelin was in high density schools, 69% of active foraging behaviour of murres (i.e. attempted contacts, approaches) was exhibited on individual capelin, compared to 24% on low density shoals and 7% on high density schools. Similarly, more murres were observed turning, a proxy of area-restricted search behaviour, when solitary and low density capelin shoals persisted for longer durations relative to when schools persisted for longer. When murres made contact with capelin (n = 16), ~70% (n = 11) were deemed successful (i.e. resulted in ingestion or ascent with fish in bill). Unsuccessful contacts resulted from fish escaping during beak manipulations to orient the fish head-first. Capelin were 7-11 times more likely to accelerate when murres displayed active versus passive (i.e. search, travel) foraging behaviours and 5-6 times more likely to accelerate in response to murre presence when in schools relative to low density shoals or solitary individuals. Overall, these results suggest that murres may increase their foraging success within areas of high prey density by preferentially searching for and targeting solitary fish that are less responsive to predators. KEY WORDS: Seabird · Common murre · Capelin · Foraging behaviour · Schooling · Antipredator · Predator–prey Full text in pdf format PreviousNextCite this article as: Crook KA, Davoren GK (2014) Underwater behaviour of common murres foraging on capelin: influences of prey density and antipredator behaviour. Mar Ecol Prog Ser 501:279-290. https://doi.org/10.3354/meps10696 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 501. Online publication date: March 31, 2014 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2014 Inter-Research.

Relative importance of local enhancement as a search strategy for breeding seabirds: an experimental approach

The spatial and temporal distribution of prey directly influences the foraging and feed- ing behaviour of predators. To investigate predator-prey interactions through the diel cycle, we examined continuous records of diving activity by a pursuit-diving seabird, the common murre Uria aalge, in conjunction with fine-scale data on the vertical distribution of their main prey, capelin Mal- lotus villosus, off the northeast Newfoundland coast, Canada. Diurnal patterns in the diving activities of murres closely reflected changes in the vertical distribution and movements of capelin. During daylight hours, 43% of murre dives were deep (≥50 m), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; ~40 to 240 m), when 82% of capelin biomass was located within or below the CIL. At night, murres concentrated diving activity at shallower depths (94% of dives were <50 m) when 86% of capelin biomass was in the upper water column. Capelin migrated through the water column during twilight periods, moving up at dusk and down at dawn. In response, murres' diving frequency increased and diving depths were graduated, becoming shallower through dusk and deeper through dawn. Crepuscular habits indicate that capelin are more accessible during twi- light periods. In summary, though murres are constrained by commuting costs, they show exceptional behavioural flexibility in their efforts to access capelin throughout their diel vertical migration (DVM). The various trade-offs involved in such predator-prey interactions are discussed, as are the ecological consequences of the DVM pattern across trophic levels.

We examined the stomach contents of 64 male and 43 female Common Murres (Uria aalge) caught in gillnets during late chick rearing/early chick departure period in August 2002 near breeding colonies in Witless Bay, Newfoundland, Canada. Twenty-six percent of stomachs contained no food. All others contained exclusively fish, and Capelin (Mallotus villosus) was the only fish positively identified. Gravid female Capelin were found in 82% of murre stomachs with contents. The number of fish ingested by murres differed between the sexes, with a mean of 2.7 fish in male stomachs and 1.6 fish in female stomachs. Mean mass of stomach contents did not differ between the sexes (males: 27.0 g; females: 22.6 g). Overall, results were similar to Common Murre diets recorded in the 1980s, despite changes in Capelin distribution and biology. The sample of Common Murres collected as gillnet bycatch was male-biased (59%). Males may have been more vulnerable to being caught in gillnets, perhaps because they were engag...

At the Gannet Islands, Labrador, the proportion of Thick-billed Murres (Uria lomvia) that successfully reared a chick that left the colony was significantly lower (mean = 63%) than that of Common Murres (Uria aalge; 82%) in each of 3 years of study. The physical and social features of the breeding site were the only factors known to influence breeding success. For both murre species, breeding success increased with ledge width, and whereas most Common Murres bred on the widest ledges, relatively few Thick-billed Murres did so. The success of each species did not differ significantly on ledges of the same width. We suggest that Common and Thick-billed murres competed for breeding sites and that Common Murres obtained a disproportionate share of the "best" sites. This occurred because Common Murres outnumbered Thick-billed Murres by about 40:1 and because cliff habitat was in short supply. Whereas Thick-billed Murres bred only on cliffs, Common Murres were able to breed on cliffs and flat ground nearer sea level. Additional circumstantial evidence supports the idea that Common and Thick-billed murres at the Gannet Islands compete for breeding sites.

As southern species undergo northward range expansions, reports of hybridization between temperate and arctic taxa are increasing, which may have important implications for the evolution, conservation, and management of arctic species. The extent of hybridization between temperate common murres (Uria aalge) and arctic thick-billed murres (U. lomvia), seabirds in the family Alcidae, has been the subject of debate. In a previous survey of variation in mitochondrial DNA (mtDNA) in common and thick-billed murres sampled from throughout the North Pacific and low Arctic, 12 of 327 common murres (~4%) were found to possess DNA sequences characteristic of thick-billed murres. In the present study, we surveyed variation in three nuclear introns in 230 common murres and 56 thick-billed murres and report that these putative hybrids carry various combinations of intron alleles from common and thick-billed murres. Analysis using the program STRUCTURE indicated that nine of these individuals possessed high proportions of thick-billed murre intron alleles, two possessed alleles in F1 and F2 proportions, and one individual possessed predominantly common murre intron alleles. We propose that the asymmetric mtDNA introgression we observed is most likely the result of mate choice at mixed colonies based on differences in male mating behaviours. Our results highlight that hybridization between thick-billed and common murres is more prevalent than previously thought, which may have important implications for the conservation and management of arctic-dwelling thick-billed murres as common murres expand northward.

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.

The foraging ecology of breeding seabirds is largely influenced by prey availability and energy demands related to reproduction which, in combination with other factors, can affect resource specialization in space and time. In this study, we combined GPS tracking, dive behavior, and stable isotope ratios of carbon and nitrogen to examine behavioral and dietary flexibility within individuals in a breeding pursuit-diving seabird, the common murre (Uria aalge), on the northeast Newfoundland coast. We examined individual flexibility over 2 years (2016, 2017) during which the timing of arrival and availability of their primary prey, capelin (Mallotus villosus), varied. In both years, we found high within-individual variation in foraging trip and dive characteristics, coupled with low spatial overlap of foraging trips both within and among individuals, indicative of flexible behavior. Concurrently, the isotopic niche of breeding common murres showed a higher degree of dietary consistency, evidenced by similar amounts of variation within and among individuals in stable isotope ratios (δ13C, δ15N). Dietary reconstruction revealed that the proportion of different prey types in the diet varied across individuals, suggesting a degree of dietary flexibility. When capelin (prey) availability was low (in 2017), foraging trips were longer and farther from the colony, maximum dive duration decreased, and the proportion of capelin in the diet decreased. Behavioral flexibility, however, remained similar across both years, regardless of prey availability. Our results suggest that common murres can tolerate and respond to fluctuations in prey availability through flexible foraging strategies. Increased foraging distances during low capelin availability suggest increased energetic costs, and thus, an energetic threshold may be reached above which lower prey availability cannot be tolerated.

During reproduction, seabirds need to balance the demands of self‐ and offspring‐provisioning within the constraints imposed by central place foraging. To assess behavioral adjustments and tolerances to these constraints, we studied the feeding tactics and reproductive success of common murres (also known as common guillemots) Uria aalge, at their largest and most offshore colony (Funk Island) where parents travel long distances to deliver a single capelin Mallotus villosus to their chicks. We assessed changes in the distance murres traveled from the colony, their proximate foraging locations and prey size choice during two successive years in which capelin exhibited an order of magnitude decrease in density and a shift from aggregated (2004) to dispersed (2005) distributions. When capelin availability was low (2005), parental murres increased their maximum foraging distances by 35% (60 to 81 km) and delivered significantly larger capelin to chicks, as predicted by central place foraging theory. Murres preferred large (&gt;140 mm) relative to small capelin (100–140 mm) in both years, but unexpectedly this preference increased as the relative density of large capelin decreased. We conclude that single prey‐loading murres target larger capelin during long foraging trips as parents are ‘forced’ to select the best prey for their offspring. Low fledgling masses suggest also that increased foraging time when capelin is scarce may come at a cost to the chicks (i.e. fewer meals per day). Murres at this colony may be functioning near physiological limits above which further or sustained adjustments in foraging effort could compromise the life‐time reproductive success of this long‐lived seabird.

We examined distributional patterns of a pursuit-diving seabird, the common murre Uria aalge, and its fish prey, capelin Mallotus villosus, within the avian foraging range of the largest murre colony in eastern North America: Funk Island, Newfoundland. During chick-rearing, the foraging habitat was previously partitioned into: (1) a high-quality area, 45 km from the colony where energy-rich capelin schools were found, which were spatially and temporally persistent and (2) a low-quality area, 60 km from the colony where schools were composed of lower-energy capelin that were ephemeral. At the scale of the foraging range (meso-scale: 1 to 100 km), murres were highly clustered into 25% of the surveyed area, with fewer murres in the low-quality relative to the high-quality area. There were tighter associations among murre and capelin aggregations in the low-quality (1.2 ± 0.2 km) relative to the high-quality area (2.6 ± 0.4 km). This likely resulted from the divergent capelin behaviour and, thus, different foraging strategies used by murres to search for (e.g. memory vs local enhancement) and capture prey. At fine spatial scales (250 m2) within foraging areas, murres were found at lower densities (mode: 2 murres), revealing that interference competition among individuals may be important during prey capture. Modeling revealed that at &gt;50 murres per 250 m2 in the high-quality area, a murre would have a &gt;90% chance of increasing its foraging efficiency by switching to forage in the low-quality area. Overall, this scale-dependent aggregative behaviour of murres suggests that cooperative foraging among conspecifics may be important in locating prey at the scale of a foraging range, or murres may simply aggregate in areas of high prey abundance, but competitive interactions among conspecifics become important at the scale of prey capture.

Juvenile capelin, Mallotus villosus (Müller), were distributed primarily on the northern Grand Banks and secondarily along the northeast coast of Newfoundland during late summer, 1994–1999. In some years, distributions extended to coastal Labrador. Capelin were seldom observed over deep water of the northeast Newfoundland Shelf or in shallow water of the southern Grand Banks. There were large areas encompassing wide ranges of capelin densities measured by the IYGPT trawl where the acoustic system did not detect capelin. Mean growth rate of capelin from 0- to 2-group was represented by length (mm)=90.9 age (years) 0.57 (r 2 =99%). The 1-group capelin formed discrete schools that varied significantly in fish length from 65 to 110 mm (p<0.05). Spatially, the different size groups occurred at scales of <55 km and possibly <3 km in some cases. The geographic distribution of age groups was not a simple function of cohort abundance. Capelin were distributed primarily in the middle ranges of temperature (5–8°C) and zooplankton biomass (2–4 g dry weight m −2 ) sampled during the surveys. It is possible that interspecific competition limits the distributions of juvenile capelin, where juvenile Arctic cod ( Boreogadus saida ) and sandlance ( Ammodytes sp.) were abundant to the north and south respectively.

We measured the fractionation of stable nitrogen (δ15N) and carbon (δ13C) isotopes in the breast and primary feathers of 11 Common Murres (Uria aalge) maintained on a diet of capelin (Mallotus villosus). Diet–feather δ15N fractionation from delipidated capelin muscle to murre feathers was 3.6‰ ± 0.2‰ in breast feathers and 3.7‰ ± 0.2‰ in primary feathers. Fractionation of δ13C was 2.5‰ ± 0.2‰ in breast feathers and 1.9‰ ± 0.3‰ in primary feathers. Prey–feather fractionation (for delipidated, muscle-only prey samples) for nine other species of seabirds ranged from 3.0‰ to 4.6‰ for δ15N and 0.1‰ to 2.5‰ for δ13C. Studies that did not remove lipids from prey samples showed higher δ15N and δ13C fractionation, and those that used whole prey items rather than muscle tissue alone showed higher δ15N fractionation. We suggest that: (1) prey samples be delipidated to facilitate interpretation of δ13C fractionation, (2) high interstudy and interspecific variation in δ13C makes species-specific studies essen...

We examined the behavioural interactions of a mobile marine predator, the common murre Uria aalge, and its prey in a cold ocean regime in the context of shifting foraging constraints of the predator during various periods of the year. To do this, we explored the scale-dependent distribution patterns of murres in relation to their prey under 2 mobility regimes: (1) during the pre-breeding period when murres can fly and (2) during the post-breeding or moulting period when murres are flightless and are accompanied by flightless juveniles. In addition, we documented for the first time the relative contributions of different prey types in the diets of murres during moult using stable isotopic analysis. Flightless adult and juvenile murres were concentrated on a meso-scale (>100 km) in an area (nursery area) where significantly greater amounts of non-capelin, mainly crustaceans, were located in closely spaced aggregations. Accordingly, flightless murres had variable proportions of fish (capelin Mallotus villosus) and crustaceans in their diet, whereas breeding murres exclusively ate capelin. At fine- and coarse-scales (0.1 to 100 km), murres that could fly during pre-breeding tracked prey at smaller spatial scales (3 to 4 km) than flightless murres (6 to 9 km). We hypothesize that varying mobility regimes and energetic demands (activity levels) of murres during these 2 periods resulted in divergent perceptions of the foraging environment and subsequently in varying tracking scales of prey. This has important implications for the scale of study during different periods of the annual cycle of a predator, when varying constraints and trophic levels may interact to produce diverse behavioural responses to prey distribution and densities.

We examine the provisioning constraints of a pursuit‐diving seabird in a cold ocean regime by comparing the behaviour of common murres Uria aalge rearing chicks at two colonies in the Northwest Atlantic during 1998‐2000. Funk Island is the largest (340,000–400,000 breeding pairs) and most offshore (60 km) colony of common murres in eastern Canada. Seventy‐five percent of the Northwest Atlantic population of common murres breeds on this island. Great Island is one island within the Witless Bay Ecological Reserve, which is the second largest breeding aggregation (100 000 breeding pairs) and is located near‐shore (2 km). The primary forage fish species in Newfoundland waters is capelin Mallotus villosus, which spawns on or near coastal beaches during summer. Therefore, the two study colonies differ in their distance to food resources and colony size. It is within this natural context that we compare: (1) prey types and frequency of delivery (amount of prey), (2) parental time budgets, and (3) the mass and condition (mass/wing length) of fledglings at both colonies. Similarly sized female capelin (100–150 mm) were delivered to chicks at both colonies. Foraging time per day per parent, a proxy of foraging effort, was similar at both colonies (Great Island: 5.1 h; Funk Island: 5.5 h), as was the percentage of time spent with mates (Great Island: 12.3%; Funk Island: 10.9%). Foraging trips, however, were longer at Funk Island (4.1 h) than at Great Island (2.9 h). This resulted in lower feeding rates of chicks (0.17 feeds per h) and poorer condition of fledglings (2.9 g/mm) at Funk Island compared to those at Great Island (0.22 feeds per h; 3.9 g/mm). We hypothesize that provisioning efforts are constrained at Funk Island by (1) distant food resources and increased competitor density, resulting in longer foraging trip durations and (2) the time spent paired with mates at the colony, which may reflect a minimum time required to maintain breeding sites due to higher breeding densities at Funk Island compared to Great Island. Demographic consequences of this poor fledgling condition at Funk Island are unknown, but fledglings may sufficiently accelerate growth at sea due to their closer proximity to an important nursery area. If fledgling survival is compromised, however, the lower potential for growth at Funk Island will impact the entire Northwest Atlantic population of common murres.