Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

Environmental determinants of distribution and foraging behaviour of cormorants (Phalacrocorax spp.) in temperate estuarine habitats

The distribution and behaviour of cormorants in estuarine environments was examined on the central coast of New South Wales, Australia, with respect to habitat associations at different spatial scales. No consistent variation in abundance was found for four species of cormorants (great Phalacrocorax carbo, pied P. varius, little black P. sulcirostris, and little pied P. melanoleucos) with state of tide (high and low) and time of day (early, middle, and late) at five estuarine locations. Differences in abundance were found among locations that were not confounded by short-term temporal variation (i.e. time of day and tide). Differences in abundance were detected among habitats (e.g. bays, creeks, and headlands) separated by hundreds of metres to kilometres in different estuaries. Cormorants of all species were rare on the open coast and near the entrance of estuaries. Abundances of cormorants varied greatly within and among creeks, bays, and river channels. Presence of seagrass beds explained much of this variation and most of the cormorants swimming and feeding were found near seagrass. Mapping of seagrass beds and the positions of cormorants at scales of metres to hundreds of metres showed a close relationship between the presence of swimming and roosting beds and the presence of seagrass beds for P. melanoleucos and for P. carbo. We argue that cormorants make decisions to visit particular estuarine habitats, especially those with seagrass, where many types of prey (e.g. fish and crustaceans) are probably most abundant. These choices must be interpreted in the context of decisions that cormorants make on scales of hundreds to thousands of kilometres during periodic excursions to the interior of Australia. Further, environmental threats to seagrass beds could impinge on these mobile visitors to the same extent as on more permanent residents. Received: 14 February 2000 / Accepted: 17 July 2000     

Intra-annual time dynamical patterns of fire sequences observed in Patagonia (Argentina)

Temporal dynamical analysis in fire sequences recorded from 1992 to 2005 in a fire-vulnerable area of Patagonia (Argentina) was performed by using the Allan Factor statistics. Three typologies of fires were investigated: forest, steppe and urban. The obtained results show the presence of annual periodicities, superimposed to a time-scaling behaviour, which characterize the point processes of fires as fractal time processes with a rather high degree of time-clusterization of the events.     

Energy budgets,risk and foraging preferences in dark-eyed juncos (Junco hyemalis)

Summary Juncos' preferences for constant versus variable food rewards were tested in three series of aviary experiments. Deprivation and feeding rates were varied across the three treatments, but the mean of the variable reward equalled the constant reward in every experiment. When the birds gained energy faster than required to meet all 24-h costs, they preferred the constant reward. When the birds' energy intake was less than the minimally required rate, they preferred the variable reward. When energy intake just balanced average daily costs, the birds preferred the constant reward or were indifferent, and their response depended on the particular mean-variance combination presented.     

Energy expenditure for mouthbrooding in a cichlid fish

Summary Energy costs of mouthbrooding were investigated in the East African maternal mouthbrooder Pseudocrenilabrus multicolor by measuring rates of oxygen consumption. Mothers with their brood in the mouth expended 15.7% more energy than mothers without their brood and 13.8% more than starving nonreproductive controls. After subtracting the energy expenditure of the brood, the excess is reduced to 4.7% and 3.2%, respectively, a difference that is not statistically significant. By contrast, feeding nonreproductive females expended more than twice as much energy as the other groups. We conclude that mouthbrooding is a low-cost strategy profiting from investments made during the preceding nonreproductive phase. The implications for the evolution of mouthbrooding are discussed within the context of ecological constraints.     

Division of foraging labor in ants can mediate demands for food and safety

Solitary foragers can balance demands for food and safety by varying their relative use of foraging patches and their level of vigilance. Here, we investigate whether colonies of the ant, Formica perpilosa, can balance these demands by dividing labor among workers. We show that foragers collecting nectar in vegetation near their nest are smaller than are those collecting nectar at sites away from the nest. We then use performance tests to show that smaller workers are more likely to succumb to attack from conspecifics but feed on nectar more efficiently than larger workers, suggesting a size-related trade-off between risk susceptibility and harvesting ability. Because foragers that travel away from the nest are probably more likely to encounter ants from neighboring colonies, this trade-off could explain the benefits of dividing foraging labor among workers. In a laboratory experiment, we show that contact with aggressive workers results in an increase in the mean size of recruits to a foraging site: this increase was not the result of more large recruits, but rather because fewer smaller ants traveled to the site. These results suggest that workers particularly susceptible to risk avoid dangerous sites, and suggest that variation in worker size can allow colonies to exploit profitably both hazardous and resource-poor patches.Communicated by L. Sundström     

Stable isotopes identify age- and sex-specific dietary partitioning and foraging habitat segregation in southern giant petrels breeding in Antarctica and southern Patagonia

We examined the isotopic signatures (δ¹³C, δ¹⁵N) of adult body feathers from southern giant petrels Macronectes giganteus collected at two breeding colonies in Antarctica (Potter Peninsula and Cape Geddes) and one in southern Patagonia (Observatorio Island), as well as in whole blood collected from adults of both sexes at each Antarctic colonies and from chicks at Potter Peninsula. As body feather moult is a continuous process in giant petrels, feathers provide an integrated annual signal of an adult’s diets and foraging habitats. In contrast, the stable isotope values of adult and chick blood are reflective of their diets during the breeding season. We found that sex-specific dietary segregation in adults breeding in Antarctica was notable during the breeding season (blood samples) but absent when examined across the entire year (feather samples). In addition, blood stable isotope values differed between chicks and adults, indicating that adults provision their offspring with a relatively higher amount of penguin and seal prey that what they consume themselves. This finding confirms previous work that suggests that chicks are preferentially fed with prey of presumably higher nutritional value such as carrion. Finally, based on isotopic differences between major oceanographic zones in the Southern Ocean, our data indicate population-specific differences in foraging distribution, with Antarctic populations move seasonally between Antarctic and subantarctic zones, while Patagonian populations likely forage in subtropical waters and in continental shelf habitats year-round.     

Genetic determination of nectar foraging,pollen foraging,and nest-site scouting in honey bee colonies

Summary Allozyme analyses of honey bee workers revealed significant differences in the intracolonial subfamily composition of groups of nectar foragers, pollen foragers, and nest-site scouts. These differences demonstrate that colony genetic structure influences the division of labor among older foraging-age bees just as it does for younger workers. The maintenance of genetic variability for the behavior of individual workers and its possible effects on the organization of colonies are discussed.     

Brooding and development of <Emphasis Type=Italic>Anasterias minuta</Emphasis> (Asteroidea: Forcipulata) in Patagonia,Argentina

Brooding, embryonic and larval development, and the influence of environmental and biological factors in tidepool habitats were studied in the sea star, Anasterias minuta, at various sites along ~220 km of the Patagonian coast. This species has a benthic, lecithotrophic development that includes eight distinct developmental stages. A larval organ, the connection cord, is developed from a small preoral lobe at early stages of development and becomes larger and thinner at advanced stages. Fecundity and average egg size increased with female body size. The regression of log egg number to log sea-star size and weight at different sites had a slope significantly less than 3.0, resulting in negative allometry and indicating that brood capacity was limited in large females. Development was generally synchronous among sites, but varied within each brood at advanced stages, with more developed brooded larvae located at the periphery of the brood mass. Brooding was synchronous among various populations at different years and spatial scales, and extended over a period of 8 months. The highest proportion of brooding females occurred during May and June (austral winter). Juveniles were released mainly during September. The likelihood of finding brooding sea stars decreased with increasing sea water temperature, tidal height, and wave exposure, and increased with increasing body size. Both body size of brooding females and brooding rate were higher in the infralittoral fringe than at midlittoral levels. A revision of the current model of brooding behavior and development among forcipulate sea stars is given.     

Movement of foraging Tundra Swans explained by spatial pattern in cryptic food densities

We tested whether Tundra Swans use information on the spatial distribution of cryptic food items (below ground Sago pondweed tubers) to shape their movement paths. In a continuous environment, swans create their own food patches by digging craters, which they exploit in several feeding bouts. Series of short (<1 m) intra-patch movements alternate with longer inter-patch movements (>1 m). Tuber biomass densities showed a positive spatial auto-correlation at a short distance (<3 m), but not at a larger distance (3-8 m). Based on the spatial pattern of the food distribution (which is assumed to be pre-harvest information for the swan) and the energy costs and benefits for different food densities at various distances, we calculated the optimal length of an inter-patch movement. A swan that moves to the patch with the highest gain rate was predicted to move to the adjacent patch (at 1 m) if the food density in the current patch had been high (>25 g/m2) and to a more distant patch (at 7-8 m) if the food density in the current patch had been low (<25 g/m2). This prediction was tested by measuring the response of swans to manipulated tuber densities. In accordance with our predictions, swans moved a long distance (>3 m) from a low-density patch and a short distance (<3 m) from a high-density patch. The quantitative agreement between prediction and observation was greater for swans feeding in pairs than for solitary swans. The result of this movement strategy is that swans visit high-density patches at a higher frequency than on offer and, consequently, achieve a 38% higher long-term gain rate. Swans also take advantage of spatial variance in food abundance by regulating the time in patches, staying longer and consuming more food from rich than from poor patches. We can conclude that the shape of the foraging path is a reflection of the spatial pattern in the distribution of tuber densities and can be understood from an optimal foraging perspective.     

Energy expenditure of swimming bottlenose dolphins (Tursiops truncatus)

The aim of our investigations was to determine, via oxygen and carbon-dioxide respirometry, how much energy dolphins (Tursiops truncatus) require when swimming at different speeds. Experiments were conducted on two female bottlenose dolphins (mean mass 162 kg) in the dolphinarium in Nuremberg Zoo, Germany, between March and August 1997. Animals were stationed in a respiration chamber for a minimum of 90 s after performing a variety of activities. We measured respiration frequency and oxygen requirements during (1) resting, (2) swimming at various velocities and (3) leaping to various heights. Resting metabolic rate of our bottlenose dolphins (2.15 W kg⁻¹) was comparable to previously published data. Metabolic rate in swimming dolphins increased to 2.47 W kg⁻¹ at 2 m s⁻¹, while leaps to 2.2 and 3 m height required a power input of 3.5 and 4 W kg⁻¹, respectively. Transport costs of swimming dolphins were lowest (1.16 J kg⁻¹ m⁻¹, corresponding to 0.12 J N⁻¹ m⁻¹) at a speed of 2.5 m s⁻¹, yielding an optimal range speed of between 1.9 and 3.2 m s⁻¹ (corresponding to minimum cost of transport ±10%). Breathing rates during all experiments correlated very well with oxygen consumption (r ² > 0.89) and could be used to derive metabolic rates in unencumbered dolphins at sea. Received: 18 December 1998 / Accepted: 27 April 1999     

Reproduction and imposex in the edible snail <Emphasis Type="Italic">Adelomelon ancilla</Emphasis> from northern Patagonia,Argentina

The edible volutid snail Adelomelon ancilla occurs on the Argentine coast and may have the capacity to serve as source of support for local fisheries. However, knowledge of its reproduction is lacking. Over 3 years (from September 2000 to July 2004) we studied the oviposition at Golfo Nuevo, Patagonia by diving. Monthly collections were performed to examine gonadal stages histologically during 2002–2003. Gamete development stages corresponded with the expected period for oviposition occurring from July to November, a time of increasing day length. Oviposition also took place in March when water temperature attained a maximum of 18°C. Following the spawning period, remaining gametes were resorpted in both sexes. Males contained sperm throughout the year, parasperm and eusperm forms were found within the same acinus. Oogonia/oocytes ranged from 20 to 240 μm in diameter during oogenesis. Females commenced laying egg capsules from a shell length of 114 mm. Reproduction is compared with other fishable volutid snails from the Argentine shelf, and suggests the need to apply certain fishing restrictions if the resource begins to be utilized commercially. The imposex condition was found in snails from some shipping areas. The frequency of this condition declined with distance from the port.     

Refueling while flying: foraging bats combust food rapidly and directly to power flight

Flying vertebrates, such as bats, face exceptionally high energy costs during active flapping flight. Once airborne, energy turnover may exceed basal metabolic rate by a factor of up to 15. Here, we asked whether fuel that powers flight originates from exogenous (dietary nutrients), endogenous sources (mostly body lipids or glycogen), or a combination of both. Since most insectivorous bats fly continuously over relatively long time periods during foraging, we assumed that slowly mobilized glycogen, although suitable for supporting brief sallying flights, is inadequate to power aerial insect-hunting of bats. We hypothesized that the insect-feeding Noctilio albiventris rapidly mobilizes and combusts nutrients from insects it has just eaten instead of utilizing endogenous lipids. We used the stable carbon isotope ratio in the bats' exhaled breath (delta13C(brth)) to assess the origin of metabolized substrates of resting and flying N. albiventris in two nutritional conditions: fasted and recently fed. The breath of fasted resting bats was depleted in 13C in relation to their insect diet (delta13C(diet)), indicating the combustion of 13C depleted body lipids. In contrast to this, delta13C(brth) of bats that had recently fed closely matched delta13C(diet) in both resting and flying bats, suggesting a quick mobilization of ingested nutrients for metabolism. In contrast to most non-volant mammals, bats have evolved the ability to fuel their high energy expenditure rates through the rapid combustion of exogenous nutrients, enabling them to conquer the nocturnal niche of aerial insectivory.     

Honeybee recruitment to scented food sources: correlations between in-hive social interactions and foraging decisions

Information exchange of environmental cues facilitates decision-making processes among members of insect societies. In honeybee foraging, it is unknown how the odor cues of a resource are relayed to inactive nest mates to enable resource exploitation at specific scented sources. It is presumed that bees need to follow the dance or to be involved in trophallaxis with a successful forager to obtain the discovered floral scent. With this in mind, we evaluated the influence of food scent relayed through in-hive interactions and the subsequent food choices. Results obtained from five colonies demonstrated that bees arriving at a feeding area preferred to land at a feeder carrying the odor currently exploited by the trained forager. The bees that landed at this feeder also showed more in-hive encounters with the trained forager than the individuals that landed at the alternative scented feeder. The most frequent interactions before landing at the correct feeder were body contacts with the active forager, a behavior that involves neither dance following nor trophallaxis. In addition, a reasonable proportion of successful newcomers showed no conspicuous interactions with the active forager. Results suggest that different sources of information can be integrated inside the hive to establish an odor-rewarded association useful to direct honeybees to a feeding site. For example, simple contacts with foragers or food exchanges with non-active foragers seem to be enough to choose a feeding site that carries the same scent collected by the focal forager.     

Shoal composition, body size and foraging in sticklebacks

Individuals which deviate from the majority in groups are likely to be most vulnerable to predation. This oddity effect, by definition, is frequency dependent, eventually fading at equal frequencies of the phenotypes in a group. It has been hypothesized that the increased predation risk of odd individuals may play an important role in the formation of phenotypically uniform shoals of fish. However, recent work has indicated that individuals may experience, or value, their predation hazard differently depending on their own size in relation to that of other group members: single large fish, but not small ones, appear concerned about their oddity in a shoal. Here I show that the apparent wariness of large fish is also expressed in a frequency-dependent manner, closely conforming to what is predicted if the oddity effect is responsible for their behavior. Using foraging activity of individuals as a means to evaluate their predation risk, I demonstrate with shoals comprising 12 threespine sticklebacks (Gasterosteus aculeatus) that large fish forage least actively when in a shoal consisting of 2 large and 10 small fish. An increase in the number of large fish to 4 among 8 small individuals clearly results in an increase in their foraging activity. However, having reached an equal frequency with small fish in a shoal, large fish do not seem to change their foraging activity much even when their number in a shoal increases further. In contrast, foraging activity of small sticklebacks remains fairly constant throughout the entire range of tested shoal compositions, providing further evidence that small and large fish respond to their oddity differently. Received: 12 February 1998 / Accepted after revision: 7 May 1998     

Age and growth in <Emphasis Type="Italic">Odontocymbiola magellanica</Emphasis> (Gastropoda: Volutidae) from Golfo Nuevo,Patagonia, Argentina

Growth, age and somatic production of the benthic predator Odontocymbiola magellanica were studied in Golfo Nuevo (42°S 65°W), on the South American Atlantic shelf. Stable oxygen isotope ratios confirmed semiannual formation of internal and external shell growth marks. Mean shell length (SL) of females was 115 and 112 mm for males, while population modal shell-free wet mass (SFWM) was 62.8 g. A Gompertz growth function (SL_∞ = 200 mm, K = 0.197, t ₀ = 5.486) fitted 113 pairs of size-at-age data (12 shells) best. O. magellanica is a long-lived species, reaching up to 20 years of age. The maximum individual somatic production of 29.3 g SFWM per year is attained at 145 mm SL, which corresponds to about 12 years of age. The life span of this volutid seems to be twice compared with other large gastropods. O. magellanica is a valuable and exploitable resource regarding its large size and somatic production, but on the other hand, its slow growth, late maturity and direct development makes it extremely vulnerable to overexploitation.