Nobuo Kokubun

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

Penguin head movement detected using small accelerometers: a proxy of prey encounter rate.

SUMMARY Determining temporal and spatial variation in feeding rates is essential for understanding the relationship between habitat features and the foraging behavior of top predators. In this study we examined the utility of head movement as a proxy of prey encounter rates in medium-sized Antarctic penguins, under the presumption that the birds should move their heads actively when they encounter and peck prey. A field study of free-ranging chinstrap and gentoo penguins was conducted at King George Island, Antarctica. Head movement was recorded using small accelerometers attached to the head, with simultaneous monitoring for prey encounter or body angle. The main prey was Antarctic krill (>99% in wet mass) for both species. Penguin head movement coincided with a slow change in body angle during dives. Active head movements were extracted using a high-pass filter (5 Hz acceleration signals) and the remaining acceleration peaks (higher than a threshold acceleration of 1.0 g) were counted. The timing of head movements coincided well with images of prey taken from the back-mounted cameras: head movement was recorded within ±2.5 s of a prey image on 89.1±16.1% (N=7 trips) of images. The number of head movements varied largely among dive bouts, suggesting large temporal variations in prey encounter rates. Our results show that head movement is an effective proxy of prey encounter, and we suggest that the method will be widely applicable for a variety of predators.

The jellyfish buffet: jellyfish enhance seabird foraging opportunities by concentrating prey

High levels of jellyfish biomass have been reported in marine ecosystems around the world, but understanding of their ecological role remains in its infancy. Jellyfish are generally thought to have indirect negative impacts on higher trophic-level predators, through changes in lower trophic pathways. However, high densities of jellyfish in the water column may affect the foraging behaviour of marine predators more directly, and the effects may not always be negative. Here, we present novel observations of a diving seabird, the thick-billed murre, feeding on fish aggregating among the long tentacles of large jellyfish, by using small video loggers attached to the birds. We show that the birds encountered large jellyfish, Chrysaora melanaster, during most of their dives, commonly fed on fish associated with jellyfish, and appeared to specifically target jellyfish with a high number of fish aggregating in their tentacles, suggesting the use of jellyfish may provide significant energetic benefits to foraging murres. We conclude that jellyfish provide feeding opportunities for diving seabirds by concentrating forage fish, and that the impacts of jellyfish on marine ecosystems are more complex than previously anticipated and may be beneficial to seabirds.

Nocturnal Foraging by Red-Legged Kittiwakes, a Surface Feeding Seabird That Relies on Deep Water Prey During Reproduction.

Narrow foraging specialization may increase the vulnerability of marine predators to climate change. The red-legged kittiwake (Rissa brevirostris) is endemic to the Bering Sea and has experienced drastic population fluctuations in recent decades, presumably due to climate-driven changes in food resources. Red-legged kittiwakes are presumed to be a nocturnal surface-foraging seabird that feed almost entirely on deep water Myctophidae fishes. However, there is little empirical evidence confirming their nocturnal foraging activity during the breeding season. This study investigated the foraging behavior of red-legged kittiwakes by combining GPS tracking, accelerometry, and dietary analyses at the world’s largest breeding colony of red-legged kittiwakes on St. George I. GPS tracking of 5 individuals revealed that 82.5% of non-flight behavior (including foraging and resting) occurred over the ocean basin (bottom depth >1,000 m). Acceleration data from 4 birds showed three types of behaviors during foraging trips: (1) flight, characterized by regular wing flapping, (2) resting on water, characterized by non-active behavior, and (3) foraging, when wing flapping was irregular. The proportions of both foraging and resting behaviors were higher at night (14.1 ± 7.1% and 20.8 ± 14.3%) compared to those during the day (6.5 ± 3.0% and 1.7 ± 2.7%). The mean duration of foraging (2.4 ± 2.9 min) was shorter than that of flight between prey patches (24.2 ± 53.1 min). Dietary analyses confirmed myctophids as the dominant prey (100% by occurrence and 98.4 ± 2.4% by wet-weight). Although the sample size was limited, these results suggest that breeding red-legged kittiwakes concentrated their foraging on myctophids available at the surface during nighttime in deep water regions. We propose that the diel patterns and ephemeral nature of their foraging activity reflected the availability of myctophids. Such foraging specialization may exacerbate the vulnerability of red-legged kittiwakes to climate change in the Bering Sea.

Isotopic values of Antarctic Krill in relation to foraging habitat of penguins

Abstract Antarctic Krill Euphausia superba is a key component of the Antarctic coastal marine ecosystem. Investigations into stable isotopic values of krill in predation hotspots are important in facilitating our understanding of the feeding environments of krill in a local coastal ecosystem. In this study we investigated stable isotopic values and maturity and size composition of krill at a small spatial scale, by logging GPS tracks of five Chinstrap Pygoscelis antarcticus and seven Gentoo P. papua penguins, and analyzing their stomach contents. The study was conducted at a penguin colony on Barton Peninsula, King George Island, Antarctica. The main food item of both species was Antarctic Krill (>98% wet mass). One Chinstrap and four Gentoo penguin foraging trips were classified as “on-shelf” trips, and four Chinstrap and three Gentoo Penguin foraging trips were classified as “off-shelf” trips. Krill collected from off-shelf trips had higher &dgr; 15N (4.22±0.28‰) values than those from on-shelf trips (3.78±0.29‰). The &dgr; 13C of the krill samples did not differ between the two penguin species or between trip types. The proportion of juvenile krill taken was higher for Chinstrap (13.04±4.97%) than Gentoo penguins (3.33±2.43%). Our results suggest that the main food source of the krill in our sample originated as non benthic planktonic/suspended organic matter, and that krill in off-shelf habitat may occasionally consume higher trophic level prey compared to those in on-shelf habitats.

Group association and vocal behaviour during foraging trips in Gentoo penguins

In contrast to their terrestrial call, the offshore call of penguins during their foraging trips has been poorly studied due to the inaccessibility of the foraging site—the open ocean—to researchers. Here, we present the first description of the vocal behaviour of penguins in the open ocean and discuss the function of their vocal communication. We deployed an animal-borne camera on gentoo penguins (Pygoscelis papua) and recorded their foraging behaviour during chick guarding. From the video recordings, we collected 598 offshore calls from 10 individuals in two breeding seasons (2014–2015 and 2015–2016), and we analysed the acoustic characteristics and behavioural contexts of these calls, including diving patterns, group association events, and foraging behaviour. The offshore calls varied in their dominant frequency and length, and penguins produced calls of different lengths in succession. Group associations were observed within one minute following an offshore call in almost half of the instances (43.18%). Penguins undertook dives of shallower depths and shorter durations after producing an offshore call than those before producing an offshore call. Our findings show that penguins may use vocal communication in the ocean related with group association during foraging trips.

Diel diving behavior of breeding gentoo penguins on King George Island in Antarctica

Many marine birds dive to catch prey in water. The gentoo penguin (Pygoscelis papua), a specialized diving seabird that preys on krill and fish, is distributed from the sub-Antarctic islands to Antarctic regions. Here, we observed the diving behavior of breeding gentoo penguins on King George Island, South Shetland Islands, Antarctica. Using a time–depth recorder, we collected diving depths every second from seven gentoo breeders for 5–21 days during the chick-rearing period and analyzed their diving characteristics. Most dives occurred in shallow water, although the dive efficiency (=bottom duration time/[dive duration + post-dive surface time]) was highest at depths of 30–35 m and decreased as the penguins dove deeper. Gentoo penguins did not dive more frequently during the day than at night, but during nighttime, most dives occurred in shallow water (<20 m) and the dive efficiency was also higher at this time. As penguins repeated their foraging trips, the number of dives, depth of dives, and trip duration did not change significantly. Our results suggest that the diel dive patterns of gentoo penguins might be related to the vertical migration of krill (upward to the surface at night). In addition, we observed that gentoo penguins could perform active diving behavior even at night, possibly aided by civil twilight during the chick-rearing period in Antarctic regions.

Identification of marine Important Bird and Biodiversity Areas for penguins around the South Shetland Islands and South Orkney Islands

Abstract Aim To provide a method of analyzing penguin tracking data to identify priority at‐sea areas for seabird conservation (marine IBAs), based on pre‐existing approaches for flying seabirds but revised according to the specific ecology of Pygoscelis penguin species. Location Waters around the Antarctic Peninsula, South Shetland, and South Orkney archipelagos (FAO Subareas 48.1 and 48.2). Methods We made key improvements to the pre‐existing protocol for identifying marine IBAs that include refining the track interpolation method and revision of parameters for the kernel analysis (smoothing factor and utilization distribution) using sensitivity tests. We applied the revised method to 24 datasets of tracking data on penguins (three species, seven colonies, and three different breeding stages—incubation, brood, and crèche). Results We identified five new marine IBAs for seabirds in the study area, estimated to hold ca. 600,000 adult penguins. Main conclusions The results demonstrate the efficacy of a new method for the designation of a network of marine IBAs in Antarctic waters for penguins based on tracking data, which can contribute to an evidence‐based, precautionary, management framework for krill fisheries.

An observation of between-mates feeding behaviour in chick-guarding chinstrap penguins

Courtship feeding has not been reported for chinstrap penguins (Pygoscelis antarctica). However, we observed a male chinstrap penguin fed to his mate during the guard stage of chick. After being fed, the female regurgitated the food to her chick. Our observation suggests that the chicks’ behaviour of begging for food may be retained in adults, but it is usually restrained.