Trophodynamics of Southern Ocean pteropods on the southern Kerguelen Plateau

Pteropods are a group of small marine gastropods that are highly sensitive to multiple stressors associated with climate change. Their trophic ecology is not well studied, with most research having focused primarily on the effects of ocean acidification on their fragile, aragonite shells. Stable isotopes analysis coupled with isotope‐based Bayesian niche metrics is useful for characterizing the trophic structure of biological assemblages. These approaches have not been implemented for pteropod assemblages. We used isotope‐based Bayesian niche metrics to investigate the trophic relationships of three co‐occurring pteropod species, with distinct feeding behaviors, sampled from the Southern Kerguelen Plateau area in the Indian Sector of the Southern Ocean—a biologically and economically important but poorly studied region. Two of these species were gymnosomes (shell‐less pteropods), which are traditionally regarded as specialist predators on other pteropods, and the third species was a thecosome (shelled pteropod), which are typically generalist omnivores. For each species, we aimed to understand (a) variability and overlap among isotopic niches; and (b) whether there was a relationship between body size and trophic position. Observed isotopic niche areas were broadest for gymnosomes, especially Clione limacina antarctica, whose observed isotopic niche area was wider than expected on both δ¹³C and δ¹⁵N value axes. We also found that trophic position significantly increased with increasing body length for Spongiobranchaea australis. We found no indication of a dietary shift toward increased trophic position with increasing body size for Clio pyramidata f. sulcata. Trophic positions ranged from 2.8 to 3.5, revealing an assemblage composed of both primary and secondary consumer behaviors. This study provides a comprehensive comparative analysis on trophodynamics in Southern Ocean pteropod species, and supports previous studies using gut content, fatty acid and stable isotope analyses. Combined, our results illustrate differences in intraspecific trophic behavior that may be attributed to differential feeding strategies at species level.     

应用脂肪酸组成研究热带东太平洋同域中上层鲨鱼营养生态位分化

物种对食物资源利用方式的差异,即营养生态位分化是物种共存的先决条件之一,对种间营养生态位的比较研究有助于了解同域分布物种的共存机制。脂肪酸组成可反映生物较长时间尺度的摄食信息,对探讨物种间营养生态位分化具有重要指示作用。热带东太平洋主要栖息有8种大型中上层鲨鱼,大青鲨(Prionace glauca)、大眼长尾鲨(Alopias superciliosus)、镰状真鲨(Carcharhinus falciformis)、长鳍真鲨(Carcharhinus longimanus)、浅海长尾鲨(Alopias pelagicus)、尖吻鲭鲨(Isurus oxyrinchus)、路氏双髻鲨(Sphyrna lewini)和锤头双髻鲨(Sphyrna zygaena),通过比较其肌肉脂肪酸组成,分析种间食性差异,营养关系及营养生态位分化。结果表明,尖吻鲭鲨营养级相对较高,大青鲨相对较低。3种鼠鲨与5种真鲨存在食性差异或栖息地隔离。浅海长尾鲨与大眼长尾鲨营养生态位重叠程度较高,存在激烈的资源竞争。大青鲨与镰状真鲨生态位宽度较大,表征其对环境的可塑性较强;尖吻鲭鲨和路氏双髻鲨生态位宽度较小,表现为其食性的特化。本研究解释了脂肪酸组成分析在鲨鱼摄食研究中的潜在应用,对分析大洋性鲨鱼的营养生态位分化,资源分配方式及同域共存机制有一定的应用价值。     

Ontogenetic diet shifts of rough scad Trachurus lathami in the North Patagonian shelf (south-west Atlantic Ocean)

Rough scad Trachurus lathami is a key pelagic fish in the Argentinean continental shelf (ACS, south-west Atlantic Ocean), with recent increases in abundance. It is a main prey of fishes and marine mammals, and shares the environment with commercially relevant pelagic species (Engraulis anchoita and Scomber colias), playing an important role linking lower and upper trophic levels in the ecosystem. This study aims to determine the ontogenetic changes in the diet composition, feeding strategy, trophic niche breadth and trophic level of T. lathami in the North Patagonian Shelf (43°–45°30′S). The stomach contents of adult fish (n = 238) were analysed. The results suggest a clear ontogenetic shift in the diet at a size of ~190 mm. Smaller individuals (160–190 mm) were specialized on misidaceans, and showed the highest trophic level, while larger T. lathami (221–230 mm) consumed decapods (Peisos petrunkevitchi) and teleosts (eggs and larvae). Trophic niche breadth was higher at the medium-sized class (191–220 mm), which mainly preyed on copepods (Calanoides carinatus) and chaetognaths (Sagitta spp.), evidencing a more diverse diet and a rather generalist strategy. Updated information on the trophic ecology of T. lathami evidences its extremely plastic feeding behaviour, being able to adapt its trophic niche to the most readily available food items from the mesopelagic community.     

New insights on the trophic ecology of blue (Prionace glauca) and shortfin mako sharks (Isurus oxyrinchus) from the oceanic eastern South Pacific

The blue shark (Prionace glauca) and the shortfin mako shark (Isurus oxyrinchus) are two large and highly migratory sharks distributed in most oceans. Although they are often caught in the south Pacific Ocean long-line fisheries, their trophic ecology is poorly understood. Stable isotopes with Bayesian mixing and dependence concentration models were performed to determine the diet and trophic differences between the two species in the South-eastern Pacific Ocean. According to the mixing models, fishes are the most important prey of these sharks. Dolphin calves and remains were found in the stomachs of both species, which represents a novel finding in trophic ecology of South Pacific sharks. Intra-specific differences were found in P. glauca, but not in specimens of I. oxyrinchus. The two sharks showed a high degree of diet overlap (73%), primarily over mackerel and dolphin carcasses. Our results indicate that blue and shortfin mako sharks have a generalist feeding strategy in the eastern Pacific Ocean, with a strong preference for teleost fishes and also for dolphin carcasses. Therefore, trophic studies are useful to understand energy flow through the food web, and the trophic position of key species.     

THE PELAGIC DISTRIBUTION OF SEA-BIRDS IN THE WESTERN INDIAN OCEAN

The present paper summarises observations made during the International Indian Ocean Expedition on board the R.R.S.' Discovery, from August to November 1963 and from February to September 1964 in the Indian Ocean north of 20° S and west of 70° E. In 1963 work was carried out in the Somali Basin. In 1964 a series of transects were made over the ocean, the main purpose of which was to investigate the system of equatorial currents and the changes in the sea associated with the onset of the Southwest Monsoon in May. The Tropical Indian Ocean is briefly described. Except for local concentrations, the surface layers are poor in nutrient salts and plankton. There is a seasonal reversal of winds and surface currents in the Arabian Sea, but seasonal changes become less marked further south. Previous ornithological observations in the western Indian Ocean are mostly confined to the Arabian Sea or to the island groups. Thus, existing information on the pelagic range of pantropical species is incomplete. Observations made on each transect across the Indian Ocean in 1964 are summarised and compared with oceanographic data collected at the same time. General conclusions are not possible on the basis of so little information, though there appeared to be some relationship between the distribution of certain species and wind or current belts. The only marked discontinuity recorded was a concentration of Puffinus pacificus and Gygis alba at the northern edge of the Equatorial Counter-current at 58° E in March. Observations made on transects that were repeated before and after the onset of the Southwest Monsoon suggested that Sterna fuscata concentrates in the equatorial region as the monsoon develops. In general, both plankton and sea-birds were more abundant at 58° E than at 67¹/₄° E. An analysis of the presence or absence of sea-birds during each observation period, which lasted an average of one hour, established the difference between pelagic species and those largely restricted to within 50 miles of their breeding stations. There was no evidence of any correlation with zooplankton abundance, though birds of all species taken together were commonest where flying-fish were most abundant and S. fuscata appeared to be commonest in cool-water areas with strong winds, i.e. the Trade-wind belts. It was not possible to sample the food organisms of sea-birds quantitatively. Sea-bird observations in the Indian Ocean more than 200 miles from the continental coasts are summarized and compared with previous observations. The little information collected in the Indian Ocean agrees with previous work in the Pacific Ocean, where sea-birds are commonest in areas of convergence, and not where local upwelling and an associated concentration of plankton occur, such as on the equator. This may be due to the fact that populations of organisms on which sea-birds feed develop or concentrate a considerable time after upwelling of nutrient-rich water occurs. Since few sea-birds were seen feeding, it seems likely that available food is scarce and that much time is required to locate areas where it is abundant. Finally, evidence is presented to indicate how some sea-bird species may avoid or reduce competition by feeding at different distances offshore, or in different geographical areas.     

FAD: Fish Aggregating Device or Fish Attracting Device? A new analysis of yellowfin tuna movements around floating objects

Tropical tuna are known to associate with floating objects. Because fish are easier to detect and to catch when around these objects, fishermen have extensively deployed a large number of artificial floating objects in the tropical oceans. Although such objects are referred to as Fish Aggregating Devices (FADs), there is yet no strong evidence that fish do show an aggregative behaviour around them. The high probability of finding tuna around FADs may be the result of an aggregation process (high density of fish because fish stay for a long time around FADs) as well as an attraction process (high flow of fish through FADs). We analysed the movements of 14 yellowfin tuna, Thunnus albacares, in relation to FADs moored in the Indian and Pacific Oceans (corresponding to all the published tracking data) to determine whether the observed movement patterns resulted from an aggregation or an attraction process. Tuna appeared to be attracted by FADs. In general, they did not stay for long close to the FADs reached, and so did not aggregate there. Some FADs may nevertheless act as Fish Aggregating Devices. The possible reasons why tuna associate with floating objects are discussed in the light of these results.     

Biogeographical variation in the diet of Holarctic martens (genus Martes,Mammalia: Carnivora: Mustelidae): adaptive foraging in generalists

Aim Studies comparing feeding habits across a genus in different geographical regions or habitats can identify factors associated with adaptive feeding behaviour, linking key ecological traits between consumers and their environment. We investigated biogeographical patterns in dietary composition and trophic diversity across the genus Martes in relation to geographical range and environmental variables. We hypothesized that widely distributed opportunistic Martes species should demonstrate adaptive variations in dietary composition and trophic diversity relative to regional geographical location (e.g. latitude, elevation), environmental variation (e.g. temperature, rainfall, snow cover and primary productivity) and concomitant variation in food supply. Location Europe, Asia and North America. Methods We examined the dietary habits of martens (Martes spp.) using original data expressed as relative frequency of occurrence, and using principal components analysis to extract the main gradients in diet composition. These were then used as response variables in regression analyses, predicted from latitude or elevation. Multiple regression analyses were performed to assess the influence of food types and environmental variables on the trophic diversity index. Results A clear latitudinal gradient in dietary composition was observed. Small mammals were the primary food type, but were less abundant in the diet of martens at lower latitude and elevation. Vegetable matter and insects were consumed more frequently in southerly and/or lower‐elevation localities. Trophic diversity was lower at higher elevation, and increased with a decline in consumption of the dominant food types, i.e. rodents, fruits and insects. Trophic diversity also increased with increasing mean temperature. Main conclusions Biogeographical variations in feeding habits across the genus Martes proved to be associated with latitude, local climate (especially temperature regime) and the availability of alternative potential foods. On an extensive geographical scale, martens respond to varying food availability by adjusting their foraging strategy and thus should be considered facultative generalists. At the species level, however, different climatic variables emerged as differentially important, indicative of adaptations to local conditions. Martes species are opportunistic and flexible feeders, and thus their conservation requires informed management, mindful of how changes in environmental conditions might influence their varied food supply.     

Extraordinary capture of a Randall's snapper Randallichthys filamentosus in the temperate south‐eastern Indian Ocean and its molecular phylogenetic relationship within the Etelinae

The capture of a rarely encountered Randall's snapper Randallichthys filamentosus (female, 587 mm fork length) from the upper continental slope (c. 350 m) off the south coast of Western Australia (c. 34·5° S; 122·5° E) in January 2014 represents its first record from the temperate Indian Ocean and a southern range extension. This record suggests that spawning of this predominantly tropical species may probably be occurring in the eastern Indian Ocean, considering the extensive, and unlikely, distance the progeny would have otherwise travelled from its typical distribution in the western and central Pacific Ocean.     

Morphometric relationships for some species of elasmobranch from tropical eastern Pacific

This study proposes length relationship equations to determine total length of four species of sharks (Carcharhinus falciformis, Mustelus lunulatus, Sphyrna lewini and Carcharhinus limbatus) based on trunk lengths (TrL) and interdorsal lengths (IL) caught in Colombia and Central America Pacific Ocean. In Central American Pacific, data were recorded for C. falciformis, C. limbatus and S. lewini between 2006 and 2012. In the Colombian Pacific, data was recorded in 1994–1995 and from 2006 to 2012 for C. falciformis, S. lewini and M. lunulatus. In the Central American Pacific the IL – TL relationship was acceptable for C. falciformis (r² = .69) and C. limbatus which presented a good relationship (r² = .81) where most specimens were juveniles; the same trend was found for S. lewini (r² = .96). The TL‐TrL relationship estimated for species caught in the Colombian Pacific was significant for C. falciformis (r² = .98) and for M. lunulatus (r² = .84); however, for S. lewini this relationship was low (r² = .55). Results indicate this is a useful tool for fishery statistics and fishery management for elasmobranch species of the Eastern Tropical Pacific.     

Diel changes in copepod respiration rates

Measurements of respiration in adult copepods at optimum concentrations in a respirometer were made at different hours of day; the amperometric method was used to measure dissolved oxygen. The migratory species from the tropical Indian Ocean and from the Black Sea displayed distinct daily variations in respiration rate and considerable difference between averages recorded in day and night hours. Such vertical migrants as Pleuromamma xiphias (Giesbr.) and Calanus euxinus Hulsemann were found to increase respiration rate late in the evening and early in the morning. No statistically reliable daily differences in the estimates of respiration were observed in copepod species like Acatria clausi (Giesbr.) and Temora discaudata (Giesbr.) which do not make long-distance migrations. In these experimental conditions the respiration rate appeared to represent changes in locomotory activity resulting from the sustained natural migratory rhythm and movements in food search. Using data available from the literature, the feeding rhythm and intensity were compared with respiration rate. This suggested that trophic status of the water may indirectly, through locomotory activity, affect the respiration rate. Increased respiration rate was suggested in the Black Sea copepod species in comparison with data obtained in earlier decades. The possibility of changes in respiration caused by changed food conditions of the environment is discussed.     

Stable isotope profiles in whale shark (<Emphasis Type="Italic">Rhincodon typus</Emphasis>) suggest segregation and dissimilarities in the diet depending on sex and size

We investigated the sex- and size-related differences in the diet of whale sharks from the Arabian Sea (north-western Indian Ocean) using carbon and nitrogen stable-isotope analyses in white muscle. The samples were collected during the commercial fishing season between April and May of 2001 in Veraval (Gujarat, India). The overall isotope signature was similar to that of the pelagic-neritic zooplanktivore Ilisha melastoma, which suggests that both species are feeding on similar prey. In whale sharks, a positive relationship was found between δ¹⁵N and δ¹³C. This, together with a significant enrichment of both heavy stable isotopes with total length indicates that the contribution to the diet of small fish and/or larger zooplankton of higher trophic level increases with the movement from offshore areas to coastal areas as they grow. Gender differences in the isotopic ratios were not statistically significant, but small sample size cannot rule out completely the existence of some degree of spatial or dietary segregation between sexes.     

Fitness and community consequences of avoiding multiple predators

We investigated the fitness and community consequences of behavioural interactions with multiple predators in a four-trophic-level system. We conducted an experiment in oval flow-through artificial-stream tanks to examine the single and interactive sublethal effects of brook trout and stoneflies on the size at emergence of Baetis bicaudatus (Ephemeroptera: Baetidae), and the cascading trophic effects on algal biomass, the food resource of the mayflies. No predation was allowed in the experiment, so that all effects were mediated through predator modifications of prey behaviour. We reared trout stream Baetis larvae from just before egg development until emergence in tanks with four treatments: (1) water from a holding tank with two brook trout (trout odour), (2) no trout odour + eight stoneflies with glued mouthparts, (3) trout odour + stoneflies and (4) no trout odour or stoneflies. We ended the experiment after 3 weeks when ten male and ten female subimagos had emerged from each tank, measured the size of ten male and ten female mature nymphs (with black wing pads), and collected algal samples from rocks at six locations in each tank. To determine the mechanism responsible for sublethal and cascading effects on lower trophic levels we made day and night observations of mayfly behaviour for the first 6 days by counting mayflies drifting in the water column and visible on natural substrata in the artificial streams. Trout odour and stoneflies similarly reduced the size of male and female Baetis emerging from artificial streams, with non-additive effects of both predators. While smaller females are less fecund, a fitness cost of small male size has not been determined. The mechanism causing sublethal effects on Baetis differed between predators. While trout stream Baetis retained their nocturnal periodicity in all treatments, stoneflies increased drift dispersal of mayflies at night, and trout suppressed night-time feeding and drift of mayflies. Stoneflies had less effect on Baetis behaviour when fish odour was present. Thus, we attribute the non-additivity of effects of fish and stoneflies on mayfly growth to an interaction modification whereby trout odour reduced the impact of stoneflies on Baetis behaviour. Since stonefly activity was also reduced in the presence of fish odour, this modification may be attributed to the effect of fish odour on stonefly behaviour. Only stoneflies delayed Baetis emergence, suggesting that stoneflies had a greater sublethal effect on Baetis fitness than did trout. Delayed emergence may reduce Baetis fitness by increasing risks of predation and parasitism on larvae, and increasing competition for mates or oviposition sites among adults. Finally, algal biomass was higher in tanks with both predators than in the other three treatments. These data implicate a behavioural trophic cascade because predators were not allowed to consume prey. Therefore, differences in algal biomass were attributed to predator-induced changes in mayfly behaviour. Our study demonstrates the importance of considering multiple predators when measuring direct sublethal effects of predators on prey fitness and indirect effects on lower trophic levels. Identification of an interaction modification illustrates the value of obtaining detailed information on behavioural mechanisms as an aid to understanding the complex interactions occurring among components of ecological communities. Received: 20 March 1997 / Accepted: 29 September 1997     

Reduction of oviposition time and enhanced larval feeding: two potential benefits of aggregative oviposition for the viburnum leaf beetle

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Report of opportunistic feeding habits by Vences’ river crab Hydrothelphusa vencesi (Brachyura: Potamoidea: Potamonautidae)

Hydrothelphusa vencesi is an endemic Madagascan freshwater crab whose ecology and behaviour are largely unknown. We report here on an episode of the feeding habits of two individuals of H. vencesi on a giant fire millipede (Aphistogoniulus infernalis) and on a cicada (Malagasia inflata) in south-east Madagascar. This represents a first report on the diet of this species and an example of its opportunistic feeding behaviour, shedding some light on the possible trophic link between an aquatic and a terrestrial system.     

Increased sea level promotes coral cover on shallow reef flats in the Andaman Sea, eastern Indian Ocean

Sea level in the Indian Ocean is subject to considerable temporal and spatial variabilities. During the period 1960–2009 at Phuket, Thailand, in the NE Indian Ocean, mean sea level increased by 2.7 mm y⁻¹. Regular monitoring of coral cover on fringing reef flats at Phuket since 1979 revealed a sensitive response of this habitat to both transient sea-level depressions and sea-level elevation. Since 1987 when more frequent sampling began, coral cover was positively correlated with the mean sea level experienced over the preceding months. Changing mean sea level explained a high proportion of the observed variation in cover, with overall increasing sea levels and a lack of negative sea-level anomalies promoting cover on the outer reef flats. Concomitantly, there have been no changes in reef community structure or any apparent shifts in zonation patterns across the reef. While future benefits of continued increases in mean sea level on reef flats in the region will be constrained by the frequency and intensity of sea-level depressions associated with the Indian Ocean Dipole, and bleaching events, the overall picture for these shallow reefs is a positive one as they respond to increasing sea level and show rapid recovery from environmental disturbances.     

Adjustment of pre‐moult foraging strategies in Macaroni Penguins Eudyptes chrysolophus according to locality,sex and breeding status

The annual moult creates the highest physiological stress during a penguin's breeding‐cycle and is preceded by a period of hyperphagia at sea. Although crucial to individual survival, foraging strategies before moult have been little investigated in keystone marine consumers in the Southern Ocean. The Macaroni Penguin Eudyptes chrysolophus demonstrates how individuals may adjust their foraging strategies during this period in line with constraints such as potential intraspecific competition between localities, foraging ability between dimorphic sexes and timing at sea between breeding and non‐breeding population components. We recorded pre‐moult behaviour at sea for 22 Macaroni Penguins from Crozet and Kerguelen Islands (southern Indian Ocean) during 2009 and 2011, using light‐based geolocation and stable isotope analysis. Penguins were distributed in population‐specific oceanic areas with similar surface temperatures (3.5 °C) south of the archipelagos, where they foraged at comparable trophic levels based on stable isotopes of their blood. Bayesian ‘broken stick’ modelling with concurrent analysis of seawater temperature records from the animal‐borne devices showed that within each population, females remained 6 days longer than males in the colder waters before heading back towards their colonies. Finally, 17 other non‐breeding individuals that moulted earlier had a higher mean blood δ¹⁵N value than did post‐breeding birds, meaning that early moulters probably fed more on fish than did late moulters. Our findings of such adjustments in foraging strategies developed across locality, sex and breeding status help understanding of the species' contrasted pre‐moult biology across its range and its ecology in the non‐breeding period.     

Penilia avirostris DANA in the Indian Ocean (Cladocera)

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Seasonal occurrence and feeding habits of black marlin, <Emphasis Type="Italic">Istiompax indica</Emphasis>, around Yonaguni Island,southwestern Japan

The seasonal occurrence of black marlin, Istiompax indica, around Yonaguni Island, southwestern Japan, was analyzed. Black marlin was abundant from April to July, and the majority of landed fish were female (97%). None of these females was reproductively mature. The condition factor of females increased from April to July. Of 56 black marlin examined, 21 individuals contained 12 prey species; the most important prey species was skipjack tuna, Katsuwonus pelamis, and one neonatal silky shark, Carcharhinus falciformis, was also found. These results indicate that the occurrence of black marlin off Yonaguni Island was associated with feeding and not with spawning.     

Temporal variation in feeding behaviour and trophic ecology of the temperate hemiramphid, <Emphasis Type="BoldItalic">Hyporhamphus melanochir</Emphasis>

The feeding ecology of southern garfish, Hyporhamphus melanochir, in South Australia was investigated to determine the dietary composition and diurnal feeding patterns, especially in relation to a hypothesised relationship between prey availability and feeding patterns. Samples of H. melanochir were collected at different times of the day and night during autumn and winter, and assigned to 3-hourly intervals. The dietary composition and feeding patterns were determined from gut contents analysis. The main food items were seagrasses (Zosteraceae) and hyperbenthic crustaceans, mainly amphipods. Polychaetes and insects were also consumed. A clear diurnal trophic shift was evident: seagrass was consumed in large volumes during the day, whilst hyperbenthic invertebrates dominated the diet during the night. Plankton samples indicated that this trophic shift reflected the higher abundances of hyperbenthic invertebrates in the water column at night. Less time was spent consuming seagrass during winter, coinciding with shorter day-lengths. H. melanochir has a strong trophic association with seagrass beds, which may account for the high garfish abundance in northern Gulf St. Vincent where extensive Zosteracean seagrass beds occur. This may be useful fundamental information for future environmentally-based fishery management decisions to help ensure the sustainability of southern garfish population and the ecosystem of which it is a part.     

The first report of Aelurostrongylus falciformis in Norwegian badgers (Meles meles)

The first report of Aelurostrongylus falciformis (Schlegel 1933) in Fennoscandian badgers is described. Routine parasitological examination of nine Norwegian badgers, at the National Veterinary Institute during 2004 and 2005, identified A. falciformis in the terminal airways of five of the animals. The first stage larvae (L1) closely resembled, in size and morphology, those of Angiostrongylus vasorum (Baillet 1866). The diagnosis for both A. falciformis and A. vasorum is frequently based on the identification of L1 in faeces or sputum. The potential for misclassification of an A. falciformis infection as A. vasorum, where larval identification is the only diagnostic method used, is discussed.