Whisker isotopic signature depicts migration patterns and multi-year intra- and inter-individual foraging strategies in fur seals

The movement and dietary history of individuals can be studied using stable isotope records in archival keratinous tissues. Here, we present a chronology of temporally fine-scale data on the trophic niche of otariid seals by measuring the isotopic signature of serially sampled whiskers. Whiskers of male Antarctic fur seals breeding at the Crozet Islands showed synchronous and regular oscillations in both their δ¹³C and δ¹⁵N values that are likely to represent their annual migrations over the long term (mean 4.8 years). At the population level, male Antarctic fur seals showed substantial variation in both δ¹³C and δ¹⁵N values, occupying nearly all the ‘isotopic space’ created by the diversity of potential oceanic habitats (from high Antarctica to the subtropics) and prey (from Antarctic krill to subantarctic and subtropical mesopelagic fishes). At the individual level, whisker isotopic signatures depict a large diversity of foraging strategies. Some seals remained in either subantarctic or Antarctic waters, while the migratory cycle of most animals encompassed a wide latitudinal gradient where they fed on different prey. The isotopic signature of whiskers, therefore, revealed new multi-year foraging strategies of male Antarctic fur seals and is a powerful tool for investigating the ecological niche during cryptic stages of mammals'' life.     

Stable isotopes document the winter foraging ecology of king penguins and highlight connectivity between subantarctic and Antarctic ecosystems

The poorly known winter foraging ecology of the king penguin, a major Southern Ocean consumer, was investigated at the subantarctic Crozet Islands where the largest global population breeds. Blood δ¹³C and δ¹⁵N values were used as proxies of the birds’ foraging habitat and diet, respectively, and circulating prolactin levels helped in determining the birds’ reproductive status. Plasma prolactin concentrations showed that king penguin adults of unknown breeding status (n = 52) that were present at the colony in winter were in fact breeders and failed breeders, but were not non ‐breeders. Circulating prolactin was neither related to δ¹³C nor δ¹⁵N values, thus suggesting that both breeders and failed breeders used the same foraging habitats and fed on the same prey. Plasma and blood cell isotopic values depicted four new relevant biological features on the feeding strategies of king penguins during the critical winter period: (1) 42% of the birds foraged in the distant Antarctic Zone, but 58% fed primarily in subantarctic waters (δ¹³C), (2) they preyed upon myctophids in both zones (δ¹⁵N), (3) individuals were consistent in their foraging strategies over the winter months (δ¹³C and δ¹⁵N), and (4) a higher proportion of females (77%–80%) than males (27%–31%) favored feeding in distant Antarctic waters (δ¹³C). This study highlights trophic connectivity between subantarctic and Antarctic ecosystems and hence the key role of energy export from Antarctic waters to sustain breeding populations of subantarctic predators, including during the Austral winter.     

Speciation chronology of rockhopper penguins inferred from molecular, geological and palaeoceanographic data

Aim The Southern Ocean is split into several biogeographical provinces between convergence zones that separate watermasses of different temperatures. Recent molecular phylogenies have uncovered a strong phylogeographic structure among rockhopper penguin populations, Eudyptes chrysocome sensu lato, from different biogeographical provinces. These studies suggested a reclassification as three species in two major clades, corresponding, respectively, to warm, subtropical and cold sub‐Antarctic watermasses rather than to geographic proximity. Such a phylogeographic pattern, also observed in plants, invertebrates and fishes of the Southern Ocean, suggests that past changes in the positions of watermasses may have affected the evolutionary history of penguins. We calculated divergence times among various rockhopper penguin clades and calibrated these data with palaeomagmatic and palaeoceanographic events to generate a speciation chronology in rockhopper penguins. Location Southern Ocean. Methods Divergence times between populations were calculated using five distinct mitochondrial DNA loci, and assuming a molecular clock model as implemented in mdiv . The molecular evolution rate of rockhopper penguins was calibrated using the radiochronological age of St Paul Island and Amsterdam Island in the southern Indian Ocean. Separations within other clades were correlated with palaeoceanographic data using this calibrated rate. Results The split between the Atlantic and Indian populations of rockhopper penguins was dated as 0.25 Ma, using the date of emergence of St Paul and Amsterdam islands, and the divergence between sub‐Antarctic and subtropical rockhopper penguins was dated as c. 0.9 Ma (i.e. during the mid‐Pleistocene transition, a major change in the Earth’s climate cycles). Main conclusions The mid‐Pleistocene transition is known to have caused a major southward shift in watermasses in the Southern Ocean, thus changing the environment around the northernmost rockhopper penguin breeding sites. This ecological isolation of northernmost populations may have caused vicariant speciation, splitting the species into two major clades. After the emergence of St Paul and Amsterdam islands in the subtropical Indian Ocean 0.25 Ma, these islands were colonized by penguins from the subtropical Atlantic, 6000 km away, rather than by penguins from the sub‐Antarctic Indian Ocean, 5000 km closer.     

Where do penguins go during the inter-breeding period? Using geolocation to track the winter dispersion of the macaroni penguin

Although penguins are key marine predators from the Southern Ocean, their migratory behaviour during the inter-nesting period remains widely unknown. Here, we report for the first time, to our knowledge, the winter foraging movements and feeding habits of a penguin species by using geolocation sensors fitted on penguins with a new attachment method. We focused on the macaroni penguin Eudyptes chrysolophus at Kerguelen, the single largest consumer of marine prey among all seabirds. Overall, macaroni penguins performed very long winter trips, remaining at sea during approximately six months within the limits of the Southern Ocean. They departed from Kerguelen in an eastward direction and distributed widely, over more than 3.10⁶ km². The penguins spent most of their time in a previously unrecognized foraging area, i.e. a narrow latitudinal band (47–49° S) within the central Indian Ocean (70–110° E), corresponding oceanographically to the Polar Frontal Zone. There, their blood isotopic niche indicated that macaroni penguins preyed mainly upon crustaceans, but not on Antarctic krill Euphausia superba, which does not occur at these northern latitudes. Such winter information is a crucial step for a better integrative approach for the conservation of this species whose world population is known to be declining.     

A northern rockhopper penguin unveils dispersion pathways in the Southern Ocean

We report the first record of a northern rockhopper penguin Eudyptes moseleyi on the Kerguelen Islands, Southern Indian Ocean. The penguin must have crossed the subtropical convergence to reach the island. This species was recently proved to be genetically different from the subantarctic eastern rockhopper penguin E. filholi that normally breeds on the Kerguelen Islands. The sequencing of a part of the mitochondrial control region shows that this bird may come from the population of Gough Island, 6,000 km away, in the south Atlantic Ocean. This finding confirms that the genetic isolation between these two penguin species is complete, although some individuals may sporadically disperse between the breeding sites. This first direct observation of a disperser from the Atlantic to the Indian Ocean also adds further support to a biogeographic dispersion pattern already suggested by phylogeographic patterns in other species from the Southern Ocean.     

King-size fast food for Antarctic fur seals

The study of scats of adult male Antarctic fur seals Arctocephalus gazella (AFS) revealed occasional frequent capture of penguins. Although AFS adult males have been occasionally reported to kill king penguins Aptenodytes patagonicus at the shore, here we report the first observations of at-sea predation by AFS on adult king penguins in the Crozet archipelago, southern Indian Ocean. During our 20 days survey, we observed 17 penguins attacked and either severely injured or killed and consumed. Only AFS adult males were seen catching king penguins successfully. Some adult females and sub-adults also attempted to catch penguins, either at sea or in land, and so did subantarctic fur seals Arctocephalus tropicalis individuals. Our results confirm the ability of fur seals to catch and predate large seabirds, even at sea.     

Long-term species, sexual and individual variations in foraging strategies of fur seals revealed by stable isotopes in whiskers

Background

Individual variations in the use of the species niche are an important component of diversity in trophic interactions. A challenge in testing consistency of individual foraging strategy is the repeated collection of information on the same individuals.

Methodology/Principal Findings

The foraging strategies of sympatric fur seals (Arctocephalus gazella and A. tropicalis) were examined using the stable isotope signature of serially sampled whiskers. Most whiskers exhibited synchronous δ¹³C and δ¹⁵N oscillations that correspond to the seal annual movements over the long term (up to 8 years). δ¹³C and δ¹⁵N values were spread over large ranges, with differences between species, sexes and individuals. The main segregating mechanism operates at the spatial scale. Most seals favored foraging in subantarctic waters (where the Crozet Islands are located) where they fed on myctophids. However, A. gazella dispersed in the Antarctic Zone and A. tropicalis more in the subtropics. Gender differences in annual time budget shape the seal movements. Males that do not perform any parental care exhibited large isotopic oscillations reflecting broad annual migrations, while isotopic values of females confined to a limited foraging range during lactation exhibited smaller changes. Limited inter-individual isotopic variations occurred in female seals and in male A. tropicalis. In contrast, male A. gazella showed large inter-individual variations, with some males migrating repeatedly to high-Antarctic waters where they fed on krill, thus meaning that individual specialization occurred over years.

Conclusions/Significance

Whisker isotopic signature yields unique long-term information on individual behaviour that integrates the spatial, trophic and temporal dimensions of the ecological niche. The method allows depicting the entire realized niche of the species, including some of its less well-known components such as age-, sex-, individual- and migration-related changes. It highlights intrapopulation heterogeneity in foraging strategies that could have important implications for likely demographic responses to environmental variability.     

Intrapopulation variability shaping isotope discrimination and turnover: experimental evidence in arctic foxes

Background

Tissue-specific stable isotope signatures can provide insights into the trophic ecology of consumers and their roles in food webs. Two parameters are central for making valid inferences based on stable isotopes, isotopic discrimination (difference in isotopic ratio between consumer and its diet) and turnover time (renewal process of molecules in a given tissue usually measured when half of the tissue composition has changed). We investigated simultaneously the effects of age, sex, and diet types on the variation of discrimination and half-life in nitrogen and carbon stable isotopes (δ¹⁵N and δ¹³C, respectively) in five tissues (blood cells, plasma, muscle, liver, nail, and hair) of a top predator, the arctic fox Vulpes lagopus.

Methodology/Principal Findings

We fed 40 farmed foxes (equal numbers of adults and yearlings of both sexes) with diet capturing the range of resources used by their wild counterparts. We found that, for a single species, six tissues, and three diet types, the range of discrimination values can be almost as large as what is known at the scale of the whole mammalian or avian class. Discrimination varied depending on sex, age, tissue, and diet types, ranging from 0.3‰ to 5.3‰ (mean  = 2.6‰) for δ¹⁵N and from 0.2‰ to 2.9‰ (mean  = 0.9‰) for δ¹³C. We also found an impact of population structure on δ¹⁵N half-life in blood cells. Varying across individuals, δ¹⁵N half-life in plasma (6 to 10 days) was also shorter than for δ¹³C (14 to 22 days), though δ¹⁵N and δ¹³C half-lives are usually considered as equal.

Conclusion/Significance

Overall, our multi-factorial experiment revealed that at least six levels of isotopic variations could co-occur in the same population. Our experimental analysis provides a framework for quantifying multiple sources of variation in isotopic discrimination and half-life that needs to be taken into account when designing and analysing ecological field studies.     

Carbon and Nitrogen Isotopes from Top Predator Amino Acids Reveal Rapidly Shifting Ocean Biochemistry in the Outer California Current

Climatic variation alters biochemical and ecological processes, but it is difficult both to quantify the magnitude of such changes, and to differentiate long-term shifts from inter-annual variability. Here, we simultaneously quantify decade-scale isotopic variability at the lowest and highest trophic positions in the offshore California Current System (CCS) by measuring δ¹⁵N and δ¹³C values of amino acids in a top predator, the sperm whale (Physeter macrocephalus). Using a time series of skin tissue samples as a biological archive, isotopic records from individual amino acids (AAs) can reveal the proximate factors driving a temporal decline we observed in bulk isotope values (a decline of ≥1 ‰) by decoupling changes in primary producer isotope values from those linked to the trophic position of this toothed whale. A continuous decline in baseline (i.e., primary producer) δ¹⁵N and δ¹³C values was observed from 1993 to 2005 (a decrease of ∼4‰ for δ¹⁵N source-AAs and 3‰ for δ¹³C essential-AAs), while the trophic position of whales was variable over time and it did not exhibit directional trends. The baseline δ¹⁵N and δ¹³C shifts suggest rapid ongoing changes in the carbon and nitrogen biogeochemical cycling in the offshore CCS, potentially occurring at faster rates than long-term shifts observed elsewhere in the Pacific. While the mechanisms forcing these biogeochemical shifts remain to be determined, our data suggest possible links to natural climate variability, and also corresponding shifts in surface nutrient availability. Our study demonstrates that isotopic analysis of individual amino acids from a top marine mammal predator can be a powerful new approach to reconstructing temporal variation in both biochemical cycling and trophic structure.     

Important marine habitat off east Antarctica revealed by two decades of multi‐species predator tracking

Satellite telemetry data are a key source of animal distribution information for marine ecosystem management and conservation activities. We used two decades of telemetry data from the East Antarctic sector of the Southern Ocean. Habitat utilization models for the spring/summer period were developed for six highly abundant, wide‐ranging meso‐ and top‐predator species: Adélie Pygoscelis adeliae and emperor Aptenodytes forsteri penguins, light‐mantled albatross Phoebetria palpebrata, Antarctic fur seals Arctocephalus gazella, southern elephant seals Mirounga leonina, and Weddell seals Leptonychotes weddellii. The regional predictions from these models were combined to identify areas utilized by multiple species, and therefore likely to be of particular ecological significance. These areas were distributed across the longitudinal breadth of the East Antarctic sector, and were characterized by proximity to breeding colonies, both on the Antarctic continent and on subantarctic islands to the north, and by sea‐ice dynamics, particularly locations of winter polynyas. These areas of important habitat were also congruent with many of the areas reported to be showing the strongest regional trends in sea ice seasonality. The results emphasize the importance of on‐shore and sea‐ice processes to Antarctic marine ecosystems. Our study provides ocean‐basin‐scale predictions of predator habitat utilization, an assessment of contemporary habitat use against which future changes can be assessed, and is of direct relevance to current conservation planning and spatial management efforts.     

A stable isotopic investigation into the causes of decline in a sub-Antarctic predator, the rockhopper penguin Eudyptes chrysocome

The rockhopper penguin (Eudyptes chrysocome) is a conspicuous apex marine predator that has experienced marked population declines throughout most of its circumpolar breeding distribution. The cause(s) for the declines remain elusive, but the relatively large spatio‐temporal scale over which population decreases have occurred implies that ecosystem‐scale, at‐sea factors are likely to be involved. We employ stable isotope analyses of carbon (¹³C/¹²C, expressed as δ¹³C) and nitrogen (¹⁵N/¹⁴N, δ¹⁵N) in time‐series of rockhopper penguin feather samples, dating back to 1861, in order to reconstruct the species' ecological history. Specifically, we examine whether rockhopper penguin population decline has been associated with a shift towards lower primary productivity in the ecosystem in which they feed, or with a shift to a diet of lower trophic status and lower quality, and we use long‐term temperature records to evaluate whether shifts in isotope ratios are associated with annual variations in sea surface temperature. Having controlled temporally for the Suess Effect and for increases in CO₂ concentrations in seawater, we found that overall, δ¹³C signatures decreased significantly over time in rockhopper penguins from seven breeding sites, supporting the hypothesis that decreases in primary productivity, and hence, carrying capacity, for which δ¹³C signature is a proxy, have been associated with the decline of penguin populations. There was some evidence of a long‐term decline in δ¹⁵N at some sites, and strong evidence that δ¹⁵N signatures were negatively related to sea surface temperatures across sites, indicative of a shift in diet to prey of lower trophic status over time and in warm years. However, a site‐by‐site analysis revealed divergent isotopic trends among sites: five of seven sites exhibited significant temporal or temperature‐related trends in isotope signatures. This study highlights the utility of stable isotope analyses when applied over relatively long timescales to apex predators.     

Foraging behaviour of the little penguin, Eudyptula minor: initial results and assessment of instrument effect

We investigated the foraging behaviour of little penguins using a new archival electronic activity recorder which simultaneously measures speed and depth against time. We present the first integrated data of foraging behaviour of two little penguins, from which we were able to distinguish between several types of travelling and foraging behaviours. The little penguins foraged mainly within 15 m of the surface and travelled at speeds between 8 and 9 km h^-1. Using attachments ranging between 1.4 and 11.8% penguin cross-sectional area (0.1 and 6% penguin mass), and isotopic water, we also assessed the effects of carrying devices while foraging. Both water influx and metabolic rates were significantly lower in penguins carrying devices, compared to penguins foraging without devices attached. Even the relatively small attachments resulted in a decreased foraging efficiency and we suggest that there is no simple or fixed relationship between size of device and the effect on the bearer.     

Non-consumptive factors affecting foraging patterns in Antarctic penguins: a review and synthesis

Recent research has clearly shown that the fear of predation, i.e. aversion to taking risks, among mesopredators or grazers, and not merely flight from an apex predator to avoid predation, is an important aspect of ecosystem structuring. In only a few, though well-documented cases, however, has this been considered in the marine environment. Herein, we review studies that have quantified behavioral responses of Adélie penguins Pygoscelis adeliae and emperor penguins Aptenodytes forsteri to the direct presence of predators, and question why the penguins avoid entering or exiting the water at night. We also show, through literature review and new analyses of Adélie penguin diving data, that Antarctic penguins are capable of successful prey capture in the dark (defined here as <3.4 lux). Finally, we summarize extensive data on seasonal migration relative to darkness and prey availability. On the basis of our findings, we propose that penguins’ avoidance of foraging at night is due to fear of predation, and not to an inability to operate effectively in darkness. We further propose that, at polar latitudes where darkness is more a seasonal than a year-round, daily feature, this “risk aversion” affects migratory movements in both species, consistent with the “trade-off” hypothesis seen in other marine vertebrates weighing foraging success against predation risk in their choice of foraging habitat. Such non-consumptive, behavioral aspects of species interactions have yet to be considered as important in Southern Ocean food webs, but may help to explain enigmatic movement patterns and choice of foraging grounds in these penguin species.     

Tracking Cats: Problems with Placing Feline Carnivores on ��18O, ��D Isoscapes

Background

Several felids are endangered and threatened by the illegal wildlife trade. Establishing geographic origin of tissues of endangered species is thus crucial for wildlife crime investigations and effective conservation strategies. As shown in other species, stable isotope analysis of hydrogen and oxygen in hair (δD_h, δ¹⁸O_h) can be used as a tool for provenance determination. However, reliably predicting the spatial distribution of δD_h and δ¹⁸O_h requires confirmation from animal tissues of known origin and a detailed understanding of the isotopic routing of dietary nutrients into felid hair.

Methodology/Findings

We used coupled δD_h and δ¹⁸O_h measurements from the North American bobcat (Lynx rufus) and puma (Puma concolor) with precipitation-based assignment isoscapes to test the feasibility of isotopic geo-location of felidae. Hairs of felid and rabbit museum specimens from 75 sites across the United States and Canada were analyzed. Bobcat and puma lacked a significant correlation between H/O isotopes in hair and local waters, and also exhibited an isotopic decoupling of δ¹⁸O_h and δD_h. Conversely, strong δD and δ¹⁸O coupling was found for key prey, eastern cottontail rabbit (Sylvilagus floridanus; hair) and white-tailed deer (Odocoileus virginianus; collagen, bone phosphate).

Conclusions/Significance

Puma and bobcat hairs do not adhere to expected pattern of H and O isotopic variation predicted by precipitation isoscapes for North America. Thus, using bulk hair, felids cannot be placed on δ¹⁸O and δD isoscapes for use in forensic investigations. The effective application of isotopes to trace the provenance of feline carnivores is likely compromised by major controls of their diet, physiology and metabolism on hair δ¹⁸O and δD related to body water budgets. Controlled feeding experiments, combined with single amino acid isotope analysis of diets and hair, are needed to reveal mechanisms and physiological traits explaining why felid hair does not follow isotopic patterns demonstrated in many other taxa.     

A method for estimating colony sizes of Adélie penguins using remote sensing imagery

Adélie penguins (Pygoscelis adeliae) are important predators of krill (Euphausia spp.) and Antarctic silverfish (Pleuragramma antarctica) during summer, are a key indicator of the status of the Southern Ocean ecosystem, and are therefore a focal species for the Committee for the Conservation of Antarctic Marine Living Resources (CCAMLR) Ecosystem Monitoring Program. The ability to monitor the population size of species potentially affected by Southern Ocean fisheries, i.e., the Adélie penguin, is critical for effective management of those resources. However, for several reasons, direct estimates of population size are not possible in many locations around Antarctica. In this study, we combine high-resolution (0.6 m) satellite imagery with spectral analysis in a supervised classification to estimate the sizes of Adélie penguin breeding colonies along Victoria Land in the Ross Sea and on the Antarctic Peninsula. Using satellite images paired with concurrent ground counts, we fit a generalized linear mixed model with Poisson errors to predict the abundance of breeding pairs as a function of the area of current-year guano staining identified in the satellite imagery. Guano-covered area proved to be an effective proxy for the number of penguins residing within. Our model provides a robust, quantitative mechanism for estimating the breeding population size of colonies captured in imagery and identifies terrain slope as a significant component influencing apparent nesting density. While our high-resolution satellite imagery technique was developed for the Adélie penguin, these principles are directly transferrable to other colonially nesting seabirds and other species that aggregate in fixed localities.     

Evidence for a krill‐rich diet from non‐destructive analyses of penguin bone

Diet strongly influences the chemistry of vertebrate soft and hard tissues. Bird bone and eggshell mineral preserve reliable records of prey consumption, even beyond the life of the predator, and analyses of hard tissues have usefully reconstructed avian diet. Here, we assess the feasibility of a non‐destructive method for distinguishing krill‐poor from krill‐rich diets in penguins. Krill (Euphausiaceae) are fluoride‐rich, and penguins that consume krill produce fluoride‐rich bones. The chemistry of bone mineral may be elucidated using Raman spectroscopy without recourse to specialised sample preparation. Published data from the diet of six penguin species were compared to a fluoride‐informative spectral band (phosphate symmetric stretch, ν₁‐PO₄^3?) in the Raman spectra of penguin humeri. Penguins that consume abundant krill (e.g. Adélie and emperor) have ν₁‐PO₄^3?‐band positions higher than 963 cm^?1, whereas penguins that primarily eat teleost fish or cephalopods (e.g. Fiordland crested, Humboldt, little blue and yellow‐eyed) have ν₁‐PO₄^3?‐band positions lower than 963 cm^?1. A krill‐rich diet can therefore be determined from the Raman spectra of penguin bones. Raman spectroscopy could be a useful supplement to existing diet analysis techniques.     

A comprehensive isotopic investigation of habitat preferences in nonbreeding albatrosses from the Southern Ocean

Albatrosses are among the world's most endangered seabirds. Threats during the nonbreeding period have major impacts on their population dynamics, but for most species, detailed information on distribution and ecology remains essentially unknown. We used stable isotope values (δ¹³C and δ¹⁵N) in feathers to infer and compare the moulting (nonbreeding) habitats of 35 populations that include all the 20 species and subspecies (444 individuals) of albatrosses breeding within the Southern Ocean and in fringing subtropical waters. Isotopic values together with a review of available information show that the 20 albatrosses can be categorized into three groups depending on their favoured moulting grounds: 12 (60%) taxa forage primarily in warm neritic waters, six (30%) in northern oceanic waters and two (10%) in oceanic waters of the Southern Ocean. Stable isotopes indicate that habitat preferences during the nonbreeding period vary much less among different breeding populations in some species (wandering, Salvin's, grey‐headed and light‐mantled sooty albatrosses), than others (black‐browed, Indian yellow‐nosed and sooty albatrosses). The major finding of our isotopic investigation is that the great majority of albatrosses spend the nonbreeding period outside the Southern Ocean, with only three species (and in the sooty albatross, just one of the breeding populations) favouring oceanic subantarctic waters at that time. Hence, the study highlights the overwhelming importance of subtropical waters for albatrosses, where the birds are known to interact with human activities and are more likely to be negatively affected by the diverse range of fisheries operating in both neritic and oceanic waters.     

Energy requirements for growth and maintenance in macaroni and rockhopper Penguins

Summary Energy requirements for growth and maintenance of macaroni and rockhopper penguin chicks at Marion Island, southern Indian Ocean, were estimated from rates of oxygen consumption and body composition analysis. Mass-specific energy expenditures of both species increased to levels 1.5 times that of hatchlings within 14–21 days of hatching and subsequently decreased. Lipid was initially accumulated slowly but the rate of accumulation increased after 30 days of age when male parents joined the females in feeding the chicks. Lipid, however, decreased markedly after 45 days of age and was presumably metabolized. Protein was laid down throughout the growth period, but doubled its initial rate of accumulation between 22 and 35 days of age. Thereafter the rate decreased until independence. Daily energy requirements of macaroni and rockhopper penguins increased from 417 and 211 kJ d^-1, respectively, in the first week after hatching to peaks of 1540 and 1170 kJ d^-1 about halfway through the growth period before decreasing until independence. Total energy requirement for growth and maintenance was estimated to be 76200 kJ for macaroni penguins and 59400 kJ for rockhopper penguins, of which growth energy comprised 38% and 28%, respectively. Based on the energy requirements and population data, macaroni and rockhopper penguin chicks at Marion Island consume an estimated 4200 and 700 t of food, respectively, each year.     

Influence of the frontal zones on ichthyoplankton and mesopelagic fish assemblages in the Crozet Basin (Indian sector of the Southern Ocean)

One of the aims of oceanographic campaign MD 68/SUZIL, carried out in austral autumn 1991 in the Indian sector of the Southern Ocean and its adjacent subtropical waters, was to investigate the influence of hydrography on the ichthyoplankton and mesopelagic fish assemblages in the Crozet Basin. It appears that, in contrast to other sectors of the Southern Ocean, the main biogeographical barriers are the Subantarctic Front and the Agulhas Front which appear to be vertical convergence fronts. The importance of the Antarctic Polar Front and the Subtropical Front as barriers to fish seems to be minimized in this area because of its particular hydrological features, such as the lack of a subantarctic zone, the maximum current intensity of the Subantarctic Front between these fronts, and their structures — they are horizontal convergence fronts.