Horizontal niche partitioning of humpback and fin whales around the West Antarctic Peninsula: evidence from a concurrent whale and krill survey

A dedicated aerial cetacean survey was conducted concurrently to a standardised net trawl survey for krill in order to investigate distribution patterns of large whales and different krill species and to investigate relationships of these. Distance sampling data were used to produce density surface models for humpback (Megaptera novaeangliae) and fin whales (Balaenoptera physalus) around the West Antarctic Peninsula (WAP). Abundance for both species was estimated over two strata in the Bransfield Strait and Drake Passage. Distinct distribution patterns suggest horizontal niche partitioning of the two whale species around the WAP, with fin whales aggregating at the shelf edge of the South Shetland Islands in the Drake Passage and humpback whales in the Bransfield Strait. Krill biomass estimated from the concurrent krill survey was used along with CTD data from the same expedition, bathymetric parameters and satellite data on chlorophyll-a and ice concentration to model krill distribution. Comparisons of the predicted distributions of both whale species with the predicted distributions of Euphausia superba, Euphausia crystallorophias and Thysanoessa macrura suggest a complex relationship rather than a straightforward correlation between krill and whales. However, results indicate that fin whales were feeding in an area dominated by T. macrura, while humpback whales were found in areas of higher E. superba biomass. Our results provide abundance estimates for humpback whales and, for the first time, fin whales in the WAP and contribute important information on feeding ecology and habitat use of these two species in the Southern Ocean.     

Penguins, fur seals, and fishing: prey requirements and potential competition in the South Shetland Islands, Antarctica

Antarctic and sub-Antarctic seabirds, marine mammals, and human fisheries concentrate their foraging efforts on a single species, Antarctic krill (Euphausiasuperba). Because these predators may have a significant effect on krill abundance, we estimated the energy and prey requirements of Adelie (Pygoscelisadeliae), chinstrap (Pygoscelisantarctica), and gentoo (Pygoscelispapua) penguins and female Antarctic fur seals (Arctocephalusgazella) breeding on the South Shetland Islands, Antarctica and compared these estimates with catch statistics from the Antarctic krill fishery. Published data on field metabolic rate, population size, diet, prey energy content, and metabolic efficiency were used to estimate prey requirements of these breeding, adult, land-based predators and their dependent offspring. Due to their large population size, chinstrap penguins were the most significant krill predators during the period examined, consuming an estimated 7.8 × 10⁸ kg krill, followed by Adelie penguins (3.1 × 10⁷ kg), gentoo penguins (1.2 × 10⁷ kg), and Antarctic fur seals (3.6 × 10⁶ kg). Total consumption of all land-based predators on the South Shetland Islands was estimated at 8.3 × 10⁸ kg krill. The commercial krill fishery harvest in the South Shetland Island region (1.0 × 10⁸ kg) was approximately 12% of this. Commercial harvest coincides seasonally and spatially with peak penguin and fur seal prey demands, and may affect prey availability to penguins and fur seals. This differs from the conclusions of Ichii et al. who asserted that the potential for competition between South Shetland predators and the commercial krill fishery is low. Received: 26 August 1997 / Accepted: 16 December 1997     

2018年夏秋季南设得兰群岛周边水域南极磷虾集群类型及其影响因素

南极磷虾是一种典型的集群性海洋生物,其集群特征为行为生态学研究领域的重要内容之一。南极磷虾在南设得兰群岛周围高度密集分布,然而磷虾集群形状和大小的机制解释仍存在较大的争议。基于南设得兰群岛周边水域收集的Simrad EK80声学数据,本研究利用Echoview V6.16软件,对声学数据进行了分析,并对磷虾集群特征进行了划分。通过主坐标分析(PCoA)检验了环境因素(表温和海况)以及时空因素对各类型磷虾集群产生的影响。结果表明:海况对磷虾集群影响较大,光照强度次之;块状小型集群的时空分布较广,夜间与白天均占有较高比例(>30%);小型集群更易出现在白天,而大型集群则更多出现在深夜; 2月,磷虾集群与海况及纬度显著相关; 3月,集群与时段显著相关; 4月,集群与时段及海况显著相关。     

Interannual variability in the feeding of ice fish (Notothenioidei, Channichthyidae) in the southern Scotia Arc and the Antarctic Peninsula region (CCAMLR Subareas 48.1 and 48.2)

We have studied the annual variation in food intake of three sub-Antarctic ice fish species (Champsocephalus gunnari, Chaenocephalus aceratus, and Pseudochaenichthys georgianus) and three high-Antarctic ice fish species (Chionodraco rastrospinosus, Cryodraco antarcticus, and Chaenodraco wilsoni). Stomach content analyses were conducted during bottom trawl surveys around the South Shetland Islands in 1998, 2001, 2002, 2003, and 2007, the South Orkney Islands in 1999 and 2009, and off the north-western Antarctic Peninsula in 2002, 2006, and 2006/2007 in order to obtain further insight into the amount of food of Antarctic demersal fish consume during summer. Annual variation in food intake was comparatively low within an area in the krill-feeding species C. gunnari and C. wilsoni. Food intake was much more variable, by a factor of 2 or 3 among years and areas, in larger C. aceratus and C. antarcticus, which rely heavily on fish as their dietary source. Food consumption was intermediate in the two species P. georgianus and C. rastrospinosus, which rely on both krill and fish.     

Aspects of the foraging behaviour of the Antarctic Tern Sterna vittata gaini at Harmony Point, South Shetland Islands

During January and February of 2002 and 2003, we studied the diet of the Antarctic Tern Sterna vittata gaini at two colonies in Nelson Island, South Shetland Islands, by identifying the prey fed to chicks by breeders. The fish Notothenia coriiceps was the main prey in both seasons, followed by the myctophid Electrona antarctica, Antarctic krill Euphausia superba and gammarid amphipods. The contribution of fish to the diet increased as chicks grew older. Fish and amphipods were brought to chicks during the day, whereas adults brought Antarctic krill at sunrise and sunset. Both the duration of the feeding trips and the number of trips per foraging bout varied according to the type of prey caught. Preliminary information suggests that, among other causes, the foraging strategy is strongly influenced by the predation pressure of skuas on chicks. Results are compared with the only two previous study on the diet of the Antarctic Tern at the South Shetland Islands.     

Green krill, the indicator of micro- and nano-size phytoplankton availability to krill

Size-fractionated chlorophyll-a concentrations of surface seawater were measured for pico-, nano-, and micro-size fractions (<2 μm, 2–10 μm, and >10 μm respectively) during commercial krill fishery operations in the waters north of the South Shetland Islands. The proportion of green krill (individuals discoloured due to active feeding on phytoplankton) had significant regressions with chlorophyll-a concentrations in micro- and nano-size fractions. Between these two fractions, chlorophyll-a concentration in the micro-size fraction showed the higher partial regression coefficient. This result shows the importance of phytoplankton larger than nano-phytoplankton, especially micro-phytoplankton, in terms of a phytoplanktonic food source for Antarctic krill in the natural environment. Accepted: 6 February 1999     

Observations of cetaceans in the south-east Atlantic sector of the Southern Ocean,during summer 2008

Knowledge of cetacean species composition and their distribution in the south-east Atlantic sector of the Southern Ocean is scarce. During a survey in February–March 2008, systematic whale sightings were carried out along transect lines following the 5° and 15° E meridians between 35° and 67° S. In total, 67 toothed whales and 126 baleen whales were observed. Both fin whales (four animals) and Antarctic minke whales Balaenoptera bonaerenses (three animals) in addition to 16 individuals of unidentified species were among the observed baleen whales. The dominating baleen whale species in our study was humpback whales Megaptera novaeangliae with 108 individuals observed. They occurred single or in groups up to seven individuals (N _mean = 2.5 ind) and eight of the counts were of calves. The relationship between humpback whale occurrence and environmental variables including Antarctic krill (Euphausia superba) abundance from acoustic recordings, hydrography, bathymetry and production was tested using general additive models. Only temperature increased the predictive power of the model with whale occurrence increasing with the decreasing temperature in more southern areas.     

Geographical variation in the diet of the Antarctic fur seal<Emphasis Type="Italic"> Arctocephalus gazella</Emphasis>

The diet of non-breeding male Antarctic fur seals Arctocephalus gazella was investigated at different localities of the Antarctic Peninsula (Cierva Point and Hope Bay), South Shetland Islands (Deception Island and Potter Peninsula) and the South Orkney Islands (Laurie Island), by the analysis of 438 scats collected from January to March 2000. The composition of the diet was diverse, with both pelagic and benthic-demersal prey represented in the samples. Antarctic krill Euphausia superba was the most frequent and numerous prey at all the study sites except at Cierva Point, followed by fish, penguins and cephalopods. Antarctic krill also predominated by mass, followed by either fish or penguins. Fish were the second most important prey by mass at the Antarctic Peninsula whereas penguins were the second most important prey by mass at the South Shetland and South Orkney Islands. Among fish, Pleuragramma antarcticum was the most important species in the diet of the Antarctic fur seals at the Antarctic Peninsula whereas Gymnoscopelus nicholsi predominated at the South Shetland and South Orkney Islands. The results are compared with previous studies, and the possibility of implementing monitoring studies on the distribution/abundance of myctophids and P. antarcticum based on the analysis of the diet of the Antarctic fur seal is considered.     

Seabirds encountered along return transects between South Africa and Antarctica in summer in relation to hydrological features

The first aim of our long-term study on the at-sea distribution of the upper trophic levels—seabirds and marine mammals—in polar marine ecosystems is to identify the main factors affecting their distribution: water masses and pack ice, fronts and ice edge as defined on the basis of water temperature, salinity and ice overage. In this study, seabird at-sea distribution was determined in the south-eastern Atlantic Ocean in summer along four return transects between Cape Town, South Africa, and Queen Maud Land, Antarctica: two on board icebreaking MS Ivan Papanin and two on board icebreaking RV Polarstern between December 2007 and January 2012. During a total of 1,930 half-an-hour transect counts devoted to seabird recording, 69,000 individuals were encountered, belonging to 57 species (mean: 36 individuals per count, all species and expeditions pooled). In comparison, the adjacent Weddell Sea shows a lower seabird biodiversity (30 species and  150 individuals per count) than in the area covered by this study. European Arctic seas reflect an intermediate biodiversity, with 30 species and 60 individuals per count; the major difference is observed in closed pack ice, almost empty in the Arctic but showing a very high biomass in the Antarctic. On the other hand, following the same route in different years allowed to compare results: density and abundance were found to be homogenous and reproducible between years for some species, while very important patchiness was detected for others, causing large heterogeneities and differences between expeditions.     

A Southern Ocean archipelago enhances feeding opportunities for a krill predator

Productivity in the oceans is heightened around oceanographic and bathymetric features such as fronts and islands. This can have a flow-on effect, providing increased food availability for higher trophic level species. Using data from a 5-day combined visual and acoustic survey, we examined the hypothesis that higher Antarctic krill (Euphausia superba) density provides a lucrative resource for humpback whales (Megaptera novaeangliae) at a remote Antarctic feeding area, the Balleny Islands (67^oS, 164°E). We assessed whale presence at the feeding area in relation to prey (krill), productivity and environmental variables using density surface modeling. We found stark differences between krill swarms near the islands and those in adjacent open water. Swarms were twice as dense and three times more numerous near the Balleny Islands compared to an open water pelagic environment, suggesting that the islands offered a profitable feeding opportunity. At the feeding area, whales were found in deeper and more productive waters with medium krill densities. These relationships, along with the high krill availability around the islands, may be the result of the Island Mass Effect.     

Benthic feeding by krill, Euphausia superba Dana, in coastal waters off West Antarctica and in Admiralty Bay, South Shetland Islands

Diatoms in the food of krill caught in the summer in coastal waters close to the South Shetland Islands and in the summer and three winters in Admiralty Bay, King George Island were investigated. The diatoms were used as indicators of sources of krill food. Principal component analysis divided the diatom taxa into two groups, one being pelagic diatoms and from the food of krill in coastal waters, and the other being benthic diatoms and taxa from the food of krill in Admiralty Bay. All pelagic taxa of diatoms were consumed by krill. In the summer in coastal waters, benthic diatoms in krill food originated from the sublittoral. In the summer in Admiralty Bay, besides phytoplankton, diatoms were noted from the sublittoral and littoral in the food of krill. However, in the winter mostly sublittoral and epiphytic diatoms were present in krill food. Accepted: 20 March 2000     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

Trophic interactions between brown and south polar skuas at Deception Island,Antarctica

It is broadly accepted that the brown skua (Stercorarius antarcticus lonnbergi) competitively excludes the south polar skua (S. maccormicki) from penguin colonies when breeding sympatrically, forcing the latter to feed on marine resources. The purpose of this work was to examine the diets and trophic niche breadths of each species where they co-occur and to determine the degree of overlap. To this end, we analyzed 169 pellets of brown skuas, collected in two different areas (20 individuals), and 152 of south polar skuas, collected in three different areas (18 individuals), on Deception Island, South Shetland Islands, Antarctica, during the austral summer 2000. Pellet analysis often underestimates the amount of easily digestible prey, but allows for comparisons of the relative contributions of different items in the diet. South polar skuas at our study locations consumed seven different food items and had a trophic niche breadth of 0.133 compared to brown skuas that fed on 10 different items and had a trophic niche breadth of 0.078. The niche overlap between the species was 82.1%. Penguins were the principal food source of both species, however, brown skuas fed mostly on chicks, while south polar skuas fed on adults (carcasses). The use of different age classes of penguins as a food source offers an alternative to competitive exclusion, allowing the coexistence of these species on Deception Island.     

Cycles of <Emphasis Type="Italic">Euphausia superba</Emphasis> recruitment evident in the diet of Pygoscelid penguins and net trawls in the South Shetland Islands,Antarctica

Size and sex of Antarctic krill taken from chinstrap and gentoo penguin diet were compared to those from scientific net surveys in the South Shetland Islands from 1998 to 2006 in order to evaluate penguin diet as a sampling mechanism and to look at trends in krill populations. Both penguin diet and net samples revealed a 4–5 year cycle in krill recruitment with one or two strong cohorts sustaining the population during each cycle. Penguin diet samples contained adult krill of similar lengths to those caught in nets; however, penguins rarely took juvenile krill. Penguin diet samples contained proportionately more females when the krill population was dominated by large adults at the end of the cycles; net samples showed greater proportions of males in these years. These patterns are comparable to those reported elsewhere in the region and are likely driven by the availability of different sizes and sexes of krill in relation to the colony.     

基于地理加权模型的南设得兰群岛北部南极磷虾渔场空间分布影响分析

结合南极磷虾渔业科学观察员收集的渔业数据和海洋环境数据,本研究利用地理加权回归模型(GWR),分析了具有空间属性的虾群深度和离岸距离两个因子,以及海水表温对南设得兰群岛北侧水域南极磷虾渔场空间分布的影响.结果表明:各年南极磷虾渔业单位捕捞努力量渔获量(CPUE)在空间上的分布无显著的集聚性;2010和2013年,3个因子之间存在空间自相关性(正相关),而2012和2016年则无自相关性.GWR模型结果显示,3个因子对CPUE的空间分布具有不同程度的影响,影响程度大小依次为虾群深度>离岸距离>温度.拟合结果发现,南设得兰群岛东、西两侧水域中表温对CPUE空间分布的影响与其他两个因子具有相反的趋势.虾群深度和离岸距离对CPUE的空间效应主要表现为负相关,但存在着年际和区域性差异.本研究结果可为南极磷虾渔场形成机制研究提供方法上的参考.     

Using carapace measurements to determine the sex of Antarctic krill, Euphausia superba

Krill (Euphausia superba) carapace measurements (length and width; mm) collected from plankton tows in the South Shetland Islands (SSI), Antarctica are used to test the generality of a common discriminant function developed to reconstruct krill length frequencies in Antarctic fur seal diets for the area surrounding South Georgia (SG). Total length and sex ratio of krill in the SSI were overestimated by 5.6 and 154%, respectively, when the SG allometric equations were applied to 3 years (2003–2005) of data. These errors arise and increase as a result of krill population dynamics, specifically recruitment that contributes large proportions of immature krill, misclassified as males by the SG discriminant function. We develop sex-specific regression models based on separate discriminant functions that provide significantly better discriminatory power. However, our analysis indicates that reconstructions of krill sex ratio and length composition in the ocean environment are less reliable in years when the ratio of immature to mature krill is high. For the SSI area, five out of 14 years (35.7%) surveyed (1992–2005) had proportions of immature to mature adult krill ≥ 0.50.     

Future recovery of baleen whales is imperiled by climate change

Historical harvesting pushed many whale species to the brink of extinction. Although most Southern Hemisphere populations are slowly recovering, the influence of future climate change on their recovery remains unknown. We investigate the impacts of two anthropogenic pressures—historical commercial whaling and future climate change—on populations of baleen whales (blue, fin, humpback, Antarctic minke, southern right) and their prey (krill and copepods) in the Southern Ocean. We use a climate–biological coupled “Model of Intermediate Complexity for Ecosystem Assessments” (MICE) that links krill and whale population dynamics with climate change drivers, including changes in ocean temperature, primary productivity and sea ice. Models predict negative future impacts of climate change on krill and all whale species, although the magnitude of impacts on whales differs among populations. Despite initial recovery from historical whaling, models predict concerning declines under climate change, even local extinctions by 2100, for Pacific populations of blue, fin and southern right whales, and Atlantic/Indian fin and humpback whales. Predicted declines were a consequence of reduced prey (copepods/krill) from warming and increasing interspecific competition between whale species. We model whale population recovery under an alternative scenario whereby whales adapt their migratory patterns to accommodate changing sea ice in the Antarctic and a shifting prey base. Plasticity in range size and migration was predicted to improve recovery for ice‐associated blue and minke whales. Our study highlights the need for ongoing protection to help depleted whale populations recover, as well as local management to ensure the krill prey base remains viable, but this may have limited success without immediate action to reduce emissions.     

Inter-annual variability in Chinstrap penguin diet at South Shetland and South Orkneys Islands

Inter-annual variability in the diet of Chinstrap penguins (Pygoscelis antarctica) at Laurie Island (South Orkney Islands) and 25 de Mayo/King George and Nelson Islands (South Shetland) was examined based on stomach contents of adults during the 2002/2003–2006/2007 and 2002/2003–2004/2005 breeding seasons, respectively. Krill (Euphausia superba) dominated the diet as frequency of occurrence (in 100% of samples), number (>99%), and percentage contribution in weight (>94.8%). Other prey items were minor and varied between years. The weight of stomach contents was significantly different. The percentage in weight of whole krill was used to compare the feeding conditions across seasons. It differed significantly at the three sites studied. Distribution of krill size varied among years and localities, showing different krill availability for penguins.     

Habitat use of hourglass dolphins near the South Shetland Islands,Antarctica

The hourglass dolphin Lagenorhynchus cruciger is the only regularly occurring small delphinid found south of the Antarctic Polar Front, yet little is known about its ecology and habitat use. This study uses 8 years (14 cruises) of standardized shipboard surveys during January–March (2003–2011) in southern Drake Passage near the South Shetland Islands to summarize the spatial distribution of hourglass dolphin sightings and quantify habitat use. Sighting data are linked to bathymetry (depth, slope) and distance to the average location of oceanographic features. A generalized linear model is used to examine the relationships between sightings and habitat features. Hourglass dolphins were sighted on 50% of surveys (n = 29); sightings were concentrated in February. Group size tended to be 2–6 individuals; there were only 2 sightings of larger groups, of 15 and 25 individuals. Sightings were distributed entirely within the deep pelagic waters north of the South Shetland Islands in southern Drake Passage and were closely associated with the southern boundary of the Antarctic Circumpolar Current. Information on occurrence and distribution reported in this study may be useful for refining habitat associations for hourglass dolphins at regional scales in the Southern Ocean.     

Competition between the krill fishery and penguins in the South Shetland Islands

Among the South Shetland Islands, the potential competition for krill (Euphausia superba) between the Japanese fishery and krill-eating breeding penguins was assessed. A low level of competition was apparent due to several factors. Spatial overlap between the main fishing and foraging areas was insignificant. Firstly, the large colonies of the dominant penguin (chinstrap penguins,Pygoscelis antarctica) occurred where sea ice disappears early in spring, and not necessarily where krill are abundant in summer, i.e. the area of krill fishery. Secondly, overlap between trawling depth and foraging dive depth of penguins was marginal, with the latter depth being shallower. Moreover, overlap in the size-frequency of krill caught by trawlers and those captured by penguins was not complete; the penguins took larger krill on average. Finally, the present small fishery is unlikely to impact upon local krill biomass in the region. Krill biomass was estimated to be 250–1500 × 10³ tonnes within the preferred fishing areas during summer. The present catch rate by the fishery (≤13 × 10³ tonnes/half-month period) is smaller by an order of magnitude, and the fishing area does not include the main foraging areas of breeding penguins.