Collective decisions in ants when foraging under crowded conditions

In this paper we examine the effect of crowding on the selection of a path in the mass-recruiting ant Lasius niger. In our experiment, ants had to go from their nest to a food source by crossing a diamond-shaped bridge, giving the choice between two paths. Two types of bridges were used: the first had two branches of equal length but different width while the second had two branches of different length and width. Experiments at high traffic volume always ended up with the selection of the wider branch, even if it was longer. This result shows that overcrowding on the narrow branch plays an essential role in the mechanism underlying the choice of route in ants. A mathematical model was developed to evaluate the importance of two mechanisms that could account for this result. The first is based on the difference in travel duration between the two paths. The second is based on the repulsive interactions between workers making head-on encounters. The model shows that travel duration per se is not sufficient to explain path choice. Rather, it is the interplay between trail following behaviour and repulsive interactions that allows ants to choose the path that minimizes their travel time. When choosing a path ants thus prefer to trade time against energy. Our results demonstrate that any environmental constraint that alters the dynamics of trail recruitment can lead to the emergence of adaptive foraging decisions without any explicit coding of information by the foragers at the individual level.     

Temporal patterns of foraging activity in some wood warblers in relation to the availability of insect prey

Summary I monitored the temporal pattern of diurnal feeding activity in several wood warbler (Parulidae) species and concomitantly recorded the numbers of active (flying) insects in 2 willow habitats in the western United States. At one site the temporal relationship between the density of active and inactive (nonflying) insects was investigated. The diurnal patterns of insect and bird activity were inversely related and each pattern was significantly nonuniform throughout the day; the wood warblers were largely inactive during the middle of the day when insects were most active.As foliage-gleaning birds, wood warblers depend primarily on the availability of inactive (nonflying) insects that they pick from the foliage, and they appear to be limited in their foraging activity by the unavailability of such insects during midday. Interestingly, the duration of midday inactivity for a given bird species varied inversely with the proportion of time that species spent flycatching. Thus, food availability may play an important role in determining the temporal patterns of feeding activity in these insectivorous bird species.     

Sex-based differences in density-dependent sociality: an experiment with a gregarious ungulate

For animals living in natural or semi-natural settings, empirical data on how sociality changes in response to increasing population density are few, especially with respect to true conspecific density and not group size. However, insight into this line of research may be far-reaching--from understanding density dependence in sexual selection to improving models of disease transmission. Using elk (Cervus elaphus Linnaeus) held in enclosures, we conducted sex-stratified experiments to test how the frequency of dyadic pairings (interaction rate) and their quality (duration) responded to manipulations in exposure to density. Using proximity-logging radio collars we recorded when and for how long individuals shared a space within 1.4 m of each other. As predicted, males increased their interaction rate as density increased. Female interaction rates, however, increased initially as density increased but soon declined to become indistinguishable from rates at low density. Females interacted for longer periods at medium densities, whereas male interaction length clearly decreased as density increased. We highlight a sexually dichotomous, density-dependent response in sociality that has yet to be reported. In addition to furthering our understanding of sociobiology (e.g., implications of time constraints presented by density on dyadic interactions), our results have implications for managing communicable disease in gregarious species of livestock and wildlife.     

Phenylacetonitrile is a male specific repellent in gregarious desert locusts, Schistocerca gregaria

Summary. The behavioural response of desert locust individuals to adult male volatiles had been investigated in a dynamic Y-T-olfactometer. Synthetic blends of a postulated adult male-produced aggregation pheromone with or without phenylacetonitrile were not attractive to fifth-instar nymphs, young or mature adults of both sexes. Instead phenylacetonitrile had a repellent effect, which was strongest in sexually mature males. The repellent effect of phenylacetonitrile was not modified by the other components of the adult male pheromone bouquet. Even at low concentrations tested the source of the pheromone was avoided. These results are in agreement with recent studies showing that this male specific pheromone is employed by mature gregarious males in sexual behaviour and acts as a courtship inhibiting pheromone. Low concentrations of phenylacetonitrile or the postulated adult male aggregation pheromone blend without phenylacetonitrile may signal the presence of swarm fellows and eventually initiate visually guided congregation behaviour in gregarious locusts. Whether this effect does significantly contribute to swarm cohesion needs further studies.     

Effects of within- and among-patch experiences on the patch-leaving decision rules in an insect parasitoid

The present study aimed to address how an insect parasitoid makes patch-departure decisions from various types of host patches and how previous patch experiences in the environment modify this decision-making process. Experiments were done with the parasitic wasp Aphidius rhopalosiphi attacking the grain aphid Sitobion avenae. In the experiments, wasps were observed in a laboratory environment containing several patches of various host densities, and behavioural records were analysed using a Coxs proportional hazards model. Consideration of the effect of the within-patch experience gave a classic pattern of patch-leaving decision rules in parasitoids: A. rhopalosiphi used local information on host quality (i.e. numbers of ovipositions or rejections) and availability (i.e. patch density) to determine departure decision. However, consideration of previous patch experiences provided evidence that these departure rules are fundamentally dynamic, responding to the physiological state of the animal and the information it has about its environment. Results showed that A. rhopalosiphi decreased its tendency to leave the visited patch after an oviposition. However, when a female has already laid several other eggs in the environment, such an incremental mechanism gradually switched to a decremental one. Hence, A. rhopalosiphi responded to egg-load depletion by leaving the visited patches sooner and by depositing a smaller number of eggs in those patches, which probably led to a decreased level of superparasitism. Results also indicated that previous experiences enabled wasps to estimate spatial host distribution and then to adjust their behavioural decisions accordingly. Thus, A. rhopalosiphi was shown to adjust its patch residence time according to the quality and the number of the patches previously visited. These proximate mechanistic rules adopted by A. rhopalosiphi females are discussed in the context of general predictions from optimality models.Communicated by D. Gwynne     

Heterogeneity in social network connections is density-dependent: implications for disease dynamics in a gregarious ungulate

Behavioral Ecology and Sociobiology - Individual animals across all taxa differ consistently in behaviour, i.e. they show personality traits. This inter-individual variability has significant...     

Male copulatory guarding enhances female foraging in a water strider

Summary In the laboratory, females of Gerris remigis foraging singly after being separated from a copulating male averaged 16 times longer to repel male copulatory attempts compared to repelling time when copulating (60 trials). In<10 min after separation, all females either became immobile at the edge (in 34% of all trials), or recopulated and either continued to forage (61%) or swam to the edge and became immobile (5%). Copulating females captured 85% of imitation prey, but single females, harassed by males, captured only 32% of prey.During a census of 2 small streams, in the pools with swimming single males, >99% of all swimming females were copulating, <1% being single. Ninety seven percent of all single females were immobile at the edge, whereas only 11% of all pairs were at the edge. In pools without swimming single males, 32% of all single females were swimming, compared to <1% in pools with swimming males. Introductions of a male into 4 pools with a female swimming singly in each resulted in the females either becoming immobile at the edge or copulating. The 2 streams had a : sex ratio of 1.6:1, and 79% of all females were copulating during the census. Foraging, copulation and copulatory attempts continued at a reduced level during the night.Thus in streams where male G. remigis are attempting to copulate, females can forage effectively only by carrying a copulating male who apparently repels copulatory attempts by other males.     

Brood neglect and contingent foraging behavior in a pelagic seabird

Among species where there is a risk to leaving offspring unattended, parents usually take alternating shifts guarding their young. However, they may occasionally exhibit brood neglect by leaving their offspring unattended at the nest. To investigate this phenomenon further, we examined the foraging behavior of the northern gannet (Morus bassanus) during chick-rearing. This species has a prolonged nestling period (13 weeks) and the single chick is usually guarded by one or other of its parents, because unattended chicks are frequently attacked by conspecifics. We tested the prediction that the foraging behavior of adults when they left their offspring alone at the nest (unattended trips) would differ in character to when adults left offspring with their partner (attended trips). Brood neglect typically occurred after a longer-than-average attendance period at the nest. Unattended trips were, on average, about half the duration of attended trips, and therefore much closer to the colony. There was also a difference in departure direction between attended and unattended trips, with unattended trips tending to be northeast of the colony. Chicks were fed by parents after both attended and unattended trips, but the frequency and the duration of unattended trips increased as chicks grew older whereas the duration of attended trips decreased as chicks grew. These results indicate that parents may be making a trade-off between risk of attack to their offspring and food-intake rate, and that the solution to this trade-off is dependent on chick age.Communicated by C. Brown     

Colony fission affects kinship in a social insect

Establishment of new groups is an important step in the life history of a social species. Fissioning is a common mode not only in group proliferation, for instance, as a regular part of the life cycle in the honey bee, but also when multiple females reproduce in the same group, as in multiple-queen ant societies. We studied the genetic consequences of fissioning in the ant Proformica longiseta, based on DNA microsatellites. In P. longiseta, new nests arise by fissioning from the old ones when they grow large, and the daughter nests consist of workers and queens or queen pupae but never both. Our results show that fissioning is not entirely random with respect to kinship. Workers tend to segregate along kin lines, but only when the initial relatedness in the parental nests is low. Workers in a daughter nest also tend to be associated with closely related adult queens, whereas such an association is not detected between workers and queen pupae. Most queens and workers are carried to the daughter nest by a specialized group of transporting workers, suggesting active kin discrimination by them. Fissioning pattern in P. longiseta is different from that found in other social insects with regular fission (e.g., the honey bee, swarm-founding wasps), where no fissioning along kin lines has been found. It does, however, resemble fissioning in another group of social animals: primates.     

Flexible foraging strategies in a diving seabird with high flight cost

How central-place foragers change search strategy in response to environmental conditions is poorly known. Foragers may vary the total distance travelled and how far they range from the central place in response to variation in the distribution of their prey. One potential reason as to why they would extend the length of their foraging trip and its distance from the colony would be to increase prey quality or quantity, despite incurring higher transit costs. To test this trade-off hypothesis in a species with high flight costs, we recorded the foraging behaviour of razorbills (Alca torca) using state-of-the-art techniques that log both individual horizontal (flight activity) and vertical (dive activity) movements. We show that the distance that razorbills travelled to foraging locations increased with sea-surface temperature, which may relate to higher prey quality or quantity. This relation is supported by an indirect index of patch quality, based on dive profiles, which also increased with travel distance from the colony. Furthermore, we show that this index was highest during the daily peak in diving activity, around midday. Taken together, these results suggest that razorbills are capable of adjusting their search strategies sensitively in response to proximate environmental cues.     

Multi-scale foraging variability in Northern gannet (Morus bassanus) fuels potential foraging plasticity

The survival of marine predators depends on behavioural plasticity to cope with changes in prey distribution. Variability in behaviour might predict plasticity and is easier to assess than plasticity. Using miniaturized GPS loggers over several breeding seasons in two Norwegian Northern gannet (Morus bassanus) colonies, we investigated if and how the variability within and between individuals, but also between colonies and years, affected foraging strategies. Results revealed strong individual variability (foraging trip durations, foraging effort and different foraging areas). Individuals from both colonies showed preferred commuting routes, flight bearings and feeding hotspots. Individuals from the largest colony used larger and more foraging areas than individuals from the small colony. Feeding hotspots and foraging ranges varied amongst years in the largest colony only. Our study demonstrated that gannets show flexibility by changing prey fields that are driven by shifting oceanographic conditions.     

Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon

Sea otter, Enhydra lutris, predation had no detectable effect on abundance and size distribution of deep-burrowing bivalve prey in the Elkhorn Slough, California, USA. Up to 23 otters were present for 6 mo of the study period (March 1984 through April 1985). This is in contrast to previous studies of sea otter predation, especially on the shallow-burrowing Pismo clam Tivela stultorum, which can be found along the wave-exposed coast near the slough. The deep-burrowing clams Tresus nuttallii and Saxidomus nuttalli made up 61% of the prey taken in the slough, and are more difficult for otters to excavate than Pismo clams. The occurrence of foraging otters was highest in an area where the two bivalve prey were extremely abundant (18 individuals m^–2). However, the otters did not selectively prey on the largest clams available within the study sight, but foraged preferentially in a patch of smaller individuals where bivalve burrow depth was restricted by the presence of a dense clay layer. This foraging strategy maximized the amount of prey biomass obtained per unit volume of sediment excavated. Our findings suggest that in soft-sediment habitats deep-burrowing bivalves may be more resistant to otter predation than shallower burrowers.     

Vigilance and foraging substrate: anti-predatory considerations in a non-standard environment

Summary The commonly studied standard anti-predatory environment presents animals with spatially-distinct feeding sites and refuges from attack, neither of which necessarily obstructs predator detection. In contrast, tree-trunks provide animals with a markedly non-standard environment in which the foraging substrate itself may be a refuge from attack that unavoidably obstructs predator detection. Thus anti-predatory behavior in this environment should be influenced not only by a perceived risk of attack, but also by the nature of the refuge/foraging substrate itself. Downy woodpeckers (Picoides pubescens) are a common tree-trunk foraging animal, and an experimental analysis of their behavior suggests that they respond appropriately to their non-standard anti-predatory environment. In particular, anti-predatory vigilance varies strongly with changes in tree trunk diameter. Two modes of vigilance were apparent. In stationary vigilance, woodpeckers maintained the position of their feet while rotating their bodies side-to-side to peer around the trunk; mobile vigilance involved movement around the trunk itself. Both the frequency and angle of rotation of stationary vigilance increased with trunk diameter, as did the frequency of mobile vigilance. The woodpeckers also held their heads farther away from the trunk surface as diameter increased. All of these measures of vigilance increased under a greater perceived risk of predation. As might be expected given these results, downy woodpeckers avoided thick trunks; they did not, however, prefer the thinnest (least obstructive) available trunks. These preferences may reflect the influence of trunk diameter on thermo-ecological and/or anti-predator considerations not related to vigilance. Overall, this arboreal environment provides an unusual perspective on anti-predator decision-making with several implications for tree-trunk foraging animals in general.     

Competition affects risk-sensitivity in foraging shrews

Summary Studies of risk-sensitive foraging have so far focused only on the effect of food demand on choice of feeding site. We suggest that competition is likely to be another factor influencing risksensitivity. A choice experiment with common shrews showed that, in the absence of competition, risk-aversion increased with increasing food intake relative to requirement. When apparent competitors were present, however, shrews were risk-indifferent regardless of their estimated requirement. The switch to risk-indifference in the presence of competitors appears to be an all-or-nothing rule of thumb which is not modified by experience with reward probability distributions.     

Tactile learning in resin foraging honeybees

Honeybees harvest and use plant resins in a mixture called propolis to seal cracks and smooth surfaces in the nest architecture. Resins in the nest may be important in maintaining a healthy colony due to their antimicrobial properties. This study had two main objectives: (1) Provide initial insight on the learning capabilities of resin foraging honeybees; (2) analyze the sensitivity of resin foraging honeybees to tactile stimuli to elucidate its possible role as a mechanism behind resin foraging. The first objective provides insight into the phenotype of these bees as compared to other forager types, while the second creates a starting point for further work on behavioral mechanisms of resin foraging. Using tactile proboscis extension response conditioning, we found that resin foragers learned to associate two different tactile stimuli, the presence of a gap between two plates and a rough sandpaper surface, with a sucrose reward significantly better than pollen foragers. The results of differential tactile conditioning exhibited no significant difference in the ability of resin foragers to discriminate between smooth and rough surfaces as compared to pollen foragers. We also determined that the sucrose response thresholds (SRTs) of returning resin foragers were lower compared to returning pollen foragers, but both resin foragers and pollen foragers learned a floral odor equally well. This is the first study to examine SRTs and conditioning to tactile and olfactory stimuli with resin foraging honeybees. The results provide new information and identify areas for future research on resin collectors, an understudied foraging phenotype.     

Chemical signals in bumble bee foraging

Summary Foraging bumblebees (Bombus vosnesenskii) deposit a substance on rewarding flowers which assists in discrimination between rewarding and nonrewarding flowers in a controlled laboratory environment. Discrimination occurs while the bee is on a flower; workers probe rewarding flowers as well as empty ones that have rewarded in the recent past, but they do not probe flowers that have had no reward. Recognition is not the result of honey contamination left on the flower by the bee during feeding. The deposit is only slightly soluble in water or ethyl alcohol but is very soluble in pentane.     

Effects of hydrolysable tannins on a herbivorous insect: fate of individual tannins in insect digestive tract

Summary. We investigated the effects of four chemically characterised galloylglucoses (GGs, a subgroup of hydrolysable tannins) and their hydrolysis product, gallic acid (GA), on consumption and performance of larvae of the autumnal moth Epirrita autumnata. Larvae were fed with birch (Betula pubescens) leaves that had been painted individually with each of the compounds at two levels, 5 and 20 mg/g. In addition, we investigated the fates of the leaf-painted GGs and GA in the E. autumnata digestive tract by comparing phenolics in leaves consumed and in faeces. In general, GGs reduced leaf consumption by E. autumnata during the second and fourth instars, although there was high compound- and instar-specific variation. However, GGs did not affect the leaf consumption rates by the most voracious fifth instar larvae. This resulted in approximately the same loss of total biomass by the experimental tree, regardless of the nature and level of GGs enriched to its foliage. The characteristic fate of hydrolysable tannins, i.e. hydrolysis, was evidenced in the larval digestive tract for three of the four leaf-painted GGs. In addition to hydrolysis, the almost total absence of GGs in larval faeces was presumably related to the oxidation of GGs. The dose-dependent excretion percentage of ingested GA showed that it's faecal content should not be used, although it commonly is, to calculate the level of GG hydrolysis. Moreover, by comparing the non-uniform appearance of faecal tetragalloylglucoses, whether ingested as such or hydrolysed from pentagalloylglucose, we concluded that a major part of oxidation of GGs occurs before their hydrolysis in the digestive tract of E. autumnata. Criticism against the common use of tannic acid, a heterogeneous mixture of GA and GGs, in ecological studies is presented. Received 15 May 2002; accepted 16 July 2002     

Jellyfish aggregations and leatherback turtle foraging patterns in a temperate coastal environment

Leatherback turtles (Dermochelys coriacea) are obligate predators of gelatinous zooplankton. However, the spatial relationship between predator and prey remains poorly understood beyond sporadic and localized reports. To examine how jellyfish (Phylum Cnidaria: Orders Semaeostomeae and Rhizostomeae) might drive the broad-scale distribution of this wide ranging species, we employed aerial surveys to map jellyfish throughout a temperate coastal shelf area bordering the northeast Atlantic. Previously unknown, consistent aggregations of Rhizostoma octopus extending over tens of square kilometers were identified in distinct coastal "hotspots" during consecutive years (2003-2005). Examination of retrospective sightings data (>50 yr) suggested that 22.5% of leatherback distribution could be explained by these hotspots, with the inference that these coastal features may be sufficiently consistent in space and time to drive long-term foraging associations.