Individual recognition of song repertoires in two wood warblers

We examined recognition of individual neighbors in hooded warblers (Wilsonia citrina) and Kentucky warblers (Oporornis formosus) by comparing responses to playbacks of neighbors' songs from appropriate and inappropriate boundaries of the subjects' territories. Kentucky warblers, which sing one song type each, responded more strongly to the songs of neighbors broadcast from incorrect than from correct boundaries. Hooded warblers use their repertoires of three to nine song types in two distinct modes of singing: repeat mode involves repeated presentation of one song type; mixed mode involves presentation of two or more song types in irregular sequence. Playbacks of neighbors' mixed-mode songs from appropriate and inappropriate boundaries indicated capabilities of individual recognition similar to those reported previously for repeat-mode songs (Godard 1991) and to those of Kentucky warblers. Repertoires of moderate size, therefore, have no pronounced influence on eventual recognition of individual neighbors in hooded warblers.     

Seasonal patterns of growth and storage in Laminaria longicruris in relation to differing patterns of availability of nitrogen in the water

The annual cycles of growth and of internal storage of nitrogen and carbon in Laminaria longicruris were compared at three sites in Nova Scotia, Canada. One site had abundant nitrogen all year round, on account of local upwelling, one had abundant nitrogen for only four months and the last was intermediate in this respect. Where nitrogen was abundant year round, growth followed the seasonal pattern of light, and there was little storage of either carbon or inorganic nitrogen. Where nitrogen was only abundant in the sea in winter, growth accelerated at this time and continued throught early summer while internal reserves of nitrogen were utilized. Carbon was stored in summer and used for growth in the fall. At the intermediate site, larger reserves of nitrogen were built up in winter and a high growth rate was maintained through most of the summer. Smaller carbon reserves were accumulated and growth was minimal in winter. Experiments in which plants were transplanted between sites were not conclusive, but suggested that plants at different sites may have genetic differences.     

Movements of intertidal whelks (Morula marginalba and Thais orbita) in relation to availability of prey and shelter

The movements of two species of predatory whelks (Muricidae: Gastropoda) were examined on several rocky seashores in New South Wales, Australia, from 1980 to 1984. The smaller species, Morula marginalba Blainville, moved less far than the larger species, Thais orbita (Gmelin). Intermittent but rapid movement allowed T. orbita to shelter in exposed places low on the shore, but to feed on their preferred prey, barnacles, much higher up to shore. This resulted in a dynamic and unpredictable pattern of vertical foraging movements for this species. Compared to areas with intact prey or with barnacles that had been killed but not removed, M. marginalba moved quickly away from areas from which barnacles had been experimentally removed. Many M. marginalba left experimental plots on two shores without prey or crevices in which to shelter, whereas some stayed over 30 d in areas with either prey or crevices (but not both), and most remained in plots where both prey and crevices were available. Thus, the presence of suitable prey and crevices seems to influence the movements of both species, which respond in different ways, corresponding to their different mobilities. These results are discussed in relation to the behavioural cues involved and predation by the whelks.     

Movement patterns of the seastar Heliaster helianthus in central Chile: relationship with environmental conditions and prey availability

The rates and patterns of feeding and displacement of predators constitute two of the most important plastic behavioral responses that allow individuals to respond quickly to changes in abundance of their prey, predation risks and to rapid alterations in environmental conditions. In this study, we quantified seasonal and spatial variation in displacement (net changes in location in 12 or 24 h periods) and prey consumed of marked individuals of the keystone seastar Heliaster helianthus at six sites spanning 600 km along the coast of north-central Chile. We evaluated the hypotheses that: (1) at sites with low availability (cover) of the main prey, the mussel Perumytilus purpuratus, Heliaster displays larger displacements and consumes a greater proportion of other prey (e.g. mobile species) than at sites with high mussel cover, (2) daily displacements will be correlated with sea surface temperature (SST) and (3) increased wave action will reduce seastar daily displacement. Our results show that Heliaster displacement is higher at sites with lower availability of P. purpuratus; and at these sites, a larger proportion of Heliaster individuals are observed feeding, mostly on other prey (e.g. limpets), which could offset the higher costs associated with increased movement. In addition, wave forces affected the activity of Heliaster negatively. Contrary to our expectations, the daily displacements did not show any relationship with SST measured on the day or the previous days of the surveys, despite the fact that average displacement was generally higher in summer than in winter months. Future studies should examine Heliaster movement during single foraging excursions and determine whether these responses affect the growth and reproductive output of individuals. Such information is vital to understand how changes in prey abundance and environmental conditions alter the behavior and energy budget of this predator and its ability to control prey populations.     

Prey vulnerability in relation to sexual coloration of prey

Sexual selection that results in the evolution of exaggerated secondary sexual characters has been hypothesized to impose production and maintenance costs of such traits on their bearers. Costs arising from sexual selection could increase the intensity of predator-mediated natural selection, leading to the prediction that species with exaggerated secondary sexual characters should be particularly susceptible to predation. We tested this prediction in a comparative analysis based on 31,745 prey individuals belonging to 66 species of birds collected from a total of 937 breeding events by 33 to 66 different pairs of European sparrowhawks Accipiter nisus annually during a period of 21 years. To assess vulnerability of different species we estimated a prey vulnerability index based on the difference in the logarithmically transformed absolute abundance of prey minus the logarithmically transformed expected abundance as determined by population density of breeding birds. The prey vulnerability index was predicted by sexual dichromatism, accounting for 23% of the variance in risk of predation among species, even when considering similarity in phenotype among species due to common descent (in the latter case explaining 12% of the variance). This finding suggests that sexual selection is an important evolutionary force-affecting predator–prey interactions.Electronic Supplementary Material Supplementary material is available for this article at     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Nectar-receiver behavior in relation to the reward rate experienced by foraging honeybees

Since forager honeybees change their food-unloading behavior according to nectar-source profitability, an experiment was performed in order to analyze whether food-receivers modify their within-hive tasks related to different reward conditions. We offered individual foragers two reward conditions at a rate feeder while an additional feeder offered a constant reward and was of free access to the rest of the hive. Both feeders were the only food sources exploited by the colony during the assays since a flight chamber was used. After receiving nectar, hive bees performed processing cycles that involved several behaviors and concluded when they returned to the delivery area to receive a new food sample. During these cycles, receivers mainly performed oral contacts offering food, or inspected cells, and often both. In the latter case, both behaviors occurred simultaneously and at the same distance from the hive entrance. When they performed a single task, either the occurrence of cell inspections increased or contact offerings decreased for the highest reward rate offered to the donor-forager. Receivers also begged for food more often after interacting with low-profit foragers. Thus, the profitability of the food source exploited by nectar-forager honeybees could affect receiver behaviors within the hives based on individual-to-individual interactions.Communicated by R.F.A. Moritz     

Temporal patterns in three-dimensional structure and activity of schools of the Atlantic silverside Menidia menidia

Temporal patterns in the structure and activity of schools of the Atlantic silverside Menidia menidia were investigated under laboratory conditions using a new computerized video technique for three-dimensional analysis. Fish were collected by seining at Middle Bridge, Pettaquamscutt River, South Kingstown, Rhode Island, USA. Approximately once a week from May to December, 1979, the behavior of a school was videotape-recorded in the laboratory at 30 min intervals from sunrise to sunset under a temperature and photoperiod regime simulating ambient field conditions. Four measurements of structure and activity were computed: (1) swimming speed (SPEED); (2) distance to nearest neighbor (NN); (3) mean direction of travel (DOT); (4) depth in the tank (DEPTH). Results demonstrated a strong seasonal rhythm in the behavior of schools, but were inconclusive in showing periodicities associated with daily, tidal or lunar cycles. NN and DEPTH were significantly different between spring, summer, fall and winter while SPEED changed significantly across 3 seasons (summer and winter were not significantly different). DOT was bimodally distributed, with peaks in spring and fall. Changes in these variables corresponded to shifts in the behavior of silversides from active, synchronized schools in spring and fall to inactive mills in summer and inactive non-schooling groups in winter. Changes in schooling behavior were only partly related to changes in temperature and photoperiod. Patterns of silverside schooling behavior correlated with their reproductive and migratory cycles and may represent an anti-predator strategy.     

Feeding rate and duration of daily activity of Astropecten aranciacus (Echinodermata: Asteroidea) in relation to prey density

In a feeding experiment, 16 individuals of Astropecten aranciacus were kept in two enclosures on the sea floor, and were provided with a surplus of food. Feeding at a daily rate of 1.3% of their dry tissue weight, their food intake was four times higher than that of free-living sea stars. The periods of daily activity were shorter than in control individuals kept in enclosures without additional food.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Temporal variability in functional responses and prey selectivity of the pelagic mysid, Mysis mixta, in natural prey assemblages

Predation rates and prey selection of the pelagic mysid shrimp, Mysis mixta, were studied experimentally in the northern Baltic Sea in 1998 during their most intensive growth period, from June to October. Functional responses during 5 months were determined by providing the mysids with a natural zooplankton assemblage, diluted to several different concentrations. The results show that ingestion rate increased, along with mysid growth, from early summer to autumn and that saturation level was reached between 400 and 500 μg C l⁻¹. Ingestion rates increased with increasing prey concentration, and sigmoidal curves explained mostly the variation in ingestion rates (explanatory levels of 86–97%). Prey selection was evident in June, July and August, though weaker during the latter 2 months. Selection differed between the studied months but, generally, copepods were more positively selected than cladocerans. Rotifers were the main prey during June and July, when mysids were small, while larger mysids fed on copepods and cladocerans. Of the copepods, Eurytemora affinis was a truly selected species. This study shows that mysids feed on many zooplankton taxa and that they undergo ontogenetic diet shifts. Received: 19 July 2000 / Accepted: 19 October 2000     

Spatial patterns of macroalgal abundance in relation to eutrophication

Based on a large monitoring data set from Danish coastal waters we tested the hypotheses: (1) The vertical pattern of algal abundance is regulated by exposure in shallow water and by light limitation towards deeper water, resulting in a bell-shaped abundance curve, with peak abundance located deeper at more exposed sites, (2) in deeper water, total algal abundance and abundance of perennial algae decrease along a eutrophication gradient while (3) abundance and relative abundance of opportunists increase. The vertical pattern of algal abundance showed a peak at intermediate water depths which was located deeper in more exposed areas and thus confirmed our first hypothesis. For algae growing from depths of peak abundance and deeper, the study demonstrated that total algal abundance and abundance of perennials and opportunists at given depths decreased significantly along a eutrophication gradient and the relationships had high explanatory power (R ² = 0.53–0.73). These results confirmed our second hypothesis. By contrast, the relative abundance of opportunists responded solely to salinity and was largest in the most brackish areas, in contradiction to hypothesis three. The lack of coupling between eutrophication and relative abundance of opportunists arises because both opportunists and the entire algal community were light limited and their ratio therefore relatively insensitive to changing water clarity. The analyses indicated that algal abundance initially responded slowly to increasing eutrophication but showed a more marked response at TN concentrations of 35–40 μM. However, the existence of possible threshold nutrient levels demands further analyses.     

How reliable are behavioral cues for assessment of male mating status in polyterritorial wood warblers,Phylloscopus sibilatrix?

It has been suggested that polyterritoriality in birds, i.e. mated males defending two spatially separated territories, deceives females into accepting polygyny against their best interest. Mated males singing on secondary territories should make it difficult for new arriving females to assess the mating status of unmated and mated males advertising to females. However, there have been objections to the deception hypothesis, suggesting that differences in territorial behavior between males of different mating status may make correct assessment possible. Polyterritorial males frequently leave their secondary territory to visit the female in the primary territory, thereby revealing their mating status to females in search of mates. The aim of this study is to investigate how reliable behavioral cues are in assessing the mating status of males in polyterritorial wood warblers. Our study shows that singing activity of mated males on secondary territories and unmated males was similar in the morning, while there was a significant difference in the afternoon. Unmated males spent, on average, more time on their territories than mated males spent on their secondary territories. However, there was considerable variation in both unmated and mated males in the frequency in which they left their territories. For instance, unmated males left their territories quite frequently early in the season compared to later. Hence, females must be able to interpret singing behavior and a male leaving his territory differently depending on time of the season and time of the day in order to correctly assess the mating status of advertising males. Estimating the increase in probability of finding an unmated male by using behavioral cues showed that females could increase the probability of finding an unmated male compared to random choice with respect to mating status. The uncertainty was, however, still considerable. The most important factor affecting the probability of finding an unmated male was the proportion of mated males on secondary territories compared to unmated males. Our study suggests that presence on a territory is not a reliable cue for the assessment of male mating status in polyterritorial wood warblers.     

A Bayesian hierarchical model of Antarctic fur seal foraging and pup growth related to sea ice and prey abundance

We created a Bayesian hierarchical model (BHM) to investigate ecosystem relationships between the physical ecosystem (sea ice extent), a prey measure (krill density), predator behaviors (diving and foraging effort of female Antarctic fur seals, Arctocephalus gazella, with pups) and predator characteristics (mass of maternal fur seals and pups). We collected data on Antarctic fur seals from 1987/1988 to 1994/1995 at Seal Island, Antarctica. The BHM allowed us to link together predators and prey into a model that uses all the data efficiently and accounts for major sources of uncertainty. Based on the literature, we made hypotheses about the relationships in the model, which we compared with the model outcome after fitting the BHM. For each BHM parameter, we calculated the mean of the posterior density and the 95% credible interval. Our model confirmed others' findings that increased sea ice was related to increased krill density. Higher krill density led to reduced dive intensity of maternal fur seals, as measured by dive depth and duration, and to less time spent foraging by maternal fur seals. Heavier maternal fur seals and lower maternal foraging effort resulted in heavier pups at 22 d. No relationship was found between krill density and maternal mass, or between maternal mass and foraging effort on pup growth rates between 22 and 85 days of age. Maternal mass may have reflected environmental conditions prior to the pup provisioning season, rather than summer prey densities. Maternal mass and foraging effort were not related to pup growth rates between 22 and 85 d, possibly indicating that food was not limiting, food sources other than krill were being used, or differences occurred before pups reached age 22 d.     

Predator foraging behavior in response to perception and learning by its prey: interactions between orb-spinning spiders and stingless bees

Although rewarded bees learn and remember colors and patterns, they have difficulty in learning to avoid negative stimuli such as decorated spider webs spun by Argiope argentata. A. argentata decorates its web with silk patterns that vary unpredictably (Fig. 1) and thus foraging insects that return to sites where spiders are found encounter new visual cues daily. Stingless bees can learn to avoid spider webs but avoidance-learning is slowed or inhibited by daily variation in web decorations (Figs. 3,4; Tables 1,2). In addition, even if bees learn to avoid decorated webs found in one location, they are unable to generalize learned-avoidance responses to similarly decorated webs found at other sites. A. argentata seems to have evolved a foraging behavior that is tied to the ways insects perceive and process information about their environment. Because of the evolutionary importance of bee-flower interdependence, the predatory behavior of web-decorating spiders may be difficult for natural selection to act against.     

Using intra-flock association patterns to understand why birds participate in mixed-species foraging flocks in terrestrial habitats

Bird species are hypothesized to join mixed-species flocks (flocks hereon) either for direct foraging or anti-predation-related benefits. In this study, conducted in a tropical evergreen forest in the Western Ghats of India, we used intra-flock association patterns to generate a community-wide assessment of flocking benefits for different species. We assumed that individuals needed to be physically proximate to particular heterospecific individuals within flocks to obtain any direct foraging benefit (flushed prey, kleptoparasitism, copying foraging locations). Alternatively, for anti-predation benefits, physical proximity to particular heterospecifics is not required, i.e. just being in the flock vicinity can suffice. Therefore, we used choice of locations within flocks to infer whether individual species are obtaining direct foraging or anti-predation benefits. A small subset of the bird community (5/29 species), composed of all members of the sallying guild, showed non-random physical proximity to heterospecifics within flocks. All preferred associates were from non-sallying guilds, suggesting that the sallying species were likely obtaining direct foraging benefits either in the form of flushed or kleptoparasitized prey. The majority of the species (24/29) chose locations randomly with respect to heterospecifics within flocks and, thus, were likely obtaining antipredation benefits. In summary, our study indicates that direct foraging benefits are important for only a small proportion of species in flocks and that predation is likely to be the main driver of flocking for most participants. Our findings apart, our study provides methodological advances that might be useful in understanding asymmetric interactions in social groups of single and multiple species.     

Correlated evolution of prey chemical discrimination with foraging, lingual morphology and vomeronasal chemoreceptor abundance in lizards

Comparative data from ten families of lizards suggest that correlated evolution has occurred between the ability to identify prey chemicals and several aspects of lingual function and morphology, abundance of vomeronasal chemoreceptor cells, and foraging behavior. Ability to discriminate prey chemicals from control substances was measured experimentally and correlated with other variables by Felsenstein's method. This ability increased with evolutionary increases in degree of lingual protrusion during tongue-flicking, which may reflect the tongue's ability to reach substrates to be sampled. It increased with deepened lingual forking and greater lingual elongation, which may be important for scent-trailing and sampling ability, respectively. Discriminatory ability also increased with abundance of vomerolfactory chemoreceptors, which presumably reflects some aspects of analytical capacities of the vomeronasal system. Prey chemical discrimination increased with degree of active foraging. Natural selection for improved vomerolfactory sampling and analysis of prey chemicals by active, but not ambush, foragers appears to account for the observed relationships. In active foragers that use vomerolfaction to locate prey, natural selection favors increased abilities to lingually sample chemicals from environmental substrates, analyze the samples for prey chemicals, and respond appropriately if prey chemicals or possible prey chemicals are detected. Such selection can account for the observed relationships among the sampling device and its movements, the sense, the discriminations, and variations in foraging ecology. Received: 13 February 1997 / Accepted after revision: 12 June 1997     

Population structure and foraging biology of the predaceous chilean asteroid Meyenaster gelatinosus and the escape biology of its prey

Observations of the feeding biology of Meyenaster gelatinosus (Meyen) were made between Horcón and the southern Golfo de Penas, Chile. Of 811 sea stars examined, 436 were feeding on individuals representing 30 prey species. M. gelatinosus preys upon almost all the echinoderms and molluscs in its habitat, yet most of the prey species have extremely effective running escape behaviour in which they eventually release their attachment to the substratum, usually assuring that they will be swept to safety. Many of the molluscs exaggerate this by dorsoventral flattening of their mantles, so that they glide even farther. The echinoid Loxechinus albus has a very effective pedicillariae defense. Even at a distance, prey species usually discern foraging M. gelatinosus from non-foraging individuals, and on several occasions were observed touching M. gelatinosus which were eating conspecifics. Comparisons of the sizes of individual M. gelatinosus and their L. albus prey items showed no correlation; none of the prey species except possibly Concholepas choncholepas and M. gelatinosus itself has a refuge in size from attacking M. gelatinosus. The density of M. gelatinosus in 3 of 4 widely separated study areas where such data were collected was 0.04/m². The mean radius of M. gelatinosus ranged from 150 to 210 mm in five study areas.     

Nitrogen availability in relation to vegetation changes resulting from grass encroachment in Dutch dry dunes

The encroachment of some tall grass species in open dune vegetation, as observed in a Dutch dry dune area, is considered unfavourable from a conservation viewpoint. This paper investigates differences in vegetation and soil properties between grass-dominated and still existing open dune grassland plots at four locations along the coast. Soil properties studied include nitrogen and phosphorus pools and nitrogen availability by mineralization. Vegetation properties included are above and below-ground biomass and nitrogen and phosphorus concentrations in above-ground biomass. Systematic differences in N-pools between grass-dominated and open dune grassland plots were not observed. However, N-availability by mineralization and its turnover rates are higher in grass-dominated plots than in open dune grassland plots, as well as above and below-ground biomass. In open dune grassland plots, atmospheric N-input is an important source of N, whereas in grass-dominated plots mineralization largely exceeds atmospheric N-input. However, these observations do not explain the mosaic-like vegetation pattern. Grazing intensity is most likely the determinant factor in the dry dune system. It is concluded, that grass encroachment is probably triggered by atmospheric deposition and is enhanced by positive feedbacks in the N-cycle. The relevance of these results for restoration management is briefly discussed.