The influence of insecticide residues on primary parasitoid and hyperparasitoid foraging behaviour in the laboratory

The behaviour of Aphidius rhopalosiphi DeStefani-Perez (Hymenoptera: Braconidae) was recorded in laboratory bioassays in the presence and absence of both residues of honeydew from the English grain aphid, Sitobion avenae (F.) and the pyrethroid insecticide, deltamethrin. Insecticide concentrations ranged from the field recommended dose rate (6.25 g ai/ha in 2001 water) to 1/16th of field rate. Parasitoids responded strongly to patches of honeydew on filter papers, however the addition of increasing concentrations of deltamethrin caused increasingly early departure from the honeydew-treated areas. Parasitoids pre-exposed to field concentration residues for between 1 and 20 min showed shorter retention times and abnormal types of behaviour on honeydew-treated patches compared to control wasps. Recovery of normal behaviour patterns occurred over a 12 h period away from the insecticide source. Further laboratory studies examined the foraging of A. rhopalosiphi and one of its associated hyperparasitoids, Dendrocerus carpenteri (Curtis), on mature wheat plants treated with an artificial honeydew solution and deltamethrin. Wasps exposed to deltamethrin residues were observed to spend shorter visit times, to groom more frequently and to rest less frequently than those on insecticide-free plants. Differences were found between the distribution of parasitoids on insecticide-treated and untreated plants, with D. carpenteri showing greater movement down insecticide-treated plants accompanied with an increase in time spent on abaxial leaf surfaces compared to A. rhopalosiphi. The results are discussed in terms of the repellency of deltamethrin and the implications for integrated pest management of differing sub-lethal insecticide effects on primary parasitoid and hyperparasitoid behaviour.     

Field study on the foraging characteristics of a ponerine ant,Hagensia havilandi Forel

Summary A field study of the foraging strategy used by the ponerine ant,Hagensia havilandi is reported. They have permanent nests in the leaf litter of coastal forests.H. havilandi is a diurnal forager and collects a variety of live and dead arthropods. These predatory ants exhibit individual foraging with no cooperation in the search for or retrieval of food items. Three colonies were observed and showed similar temporal and spatial foraging patterns. The paths of individual ants were followed and the results showed that the foragers exhibit area fidelity, and return to the nest via a direct route on finding on prey item. Several foragers did not return to the nest at dusk but returned the following morning. Occasionally a limited amount of tandem recruitment was displayed.     

Influence of host distribution on foraging behaviour in the hyperparasitoid wasp, Dendrocerus carpenteri

The foraging behaviour of Dendrocerus carpenteriCurtis (Hymenoptera: Megaspilidae), an ectophagous hyperparasitoid of aphidiine wasps inside mummified aphids, was examined in the laboratory with an experimental system consisting of broad bean, Vicia fabaL, the pea aphid, Acyrthosiphon pisumHarris, and a primary parasitoid, Ephedrus californicusBaker. Pea aphids parasitised by E. californicusoften disperse from their feeding sites (or off host plants) before dying and mummifying. Response of female hyperparasitoids to host distribution was evaluated at two spatial scales. At the first scale, behaviour of hyperparasitoids was examined on individual plants with different densities of hosts. At the second scale, habitat complexity and host location were manipulated in large foraging cages containing several plants. I show that patterns of density-dependent hyperparasitism can result from the foraging behaviour of D. carpenteri. However, dispersal of parasitised aphids may not reduce the incidence of hyperparasitism if hyperparasitoids systematically search the habitat.     

Within-patch foraging behaviour of the aphid parasitoidAphidius funebris: plant architecture, host behaviour, and individual variation

The influence of plant architecture, host colony size, and host colony structure on the foraging behaviour of the aphid parasitoidAphidius funebris Mackauer (Hymenoptera: Aphidiidae) was investigated using a factorial experimental design. The factorial design involved releasing individual parasitoid females in aphid colonies consisting of either 10 or 20 individuals ofUroleucon jaceae L. (Homoptera: Aphididae) of either only larval instar L3 or a mixture of host instars, both on unmanipulated plants and on plants that had all leaves adjacent to the colony removed. Interactions between the parasitoid and its host were recorded until the parasitoid had left the plant. The time females spent on the host plant and the number of eggs laid varied greatly among females. Host colony size significantly affected patch residence time and the number of contacts between parasitoids and aphids. Plant architecture influenced the time-budget of the parasitoids which used leaves adjacent to the aphid colony for attacking aphids. Female oviposition rate was higher on unmanipulated plants than on manipulated plants. No further significant treatment effects on patch residence time, the number of contacts, attacks or ovipositions were found. Oviposition success ofA. funebris was influenced by instar-specific host behaviour. Several rules-of-thumb proposed by foraging theory did not account for parasitoid patch-leaving behaviour.     

Field studies of foraging behaviour of an intertidal beetle

Abstract. 1. Adult staphylinid beetles Thinopinus pictus LeC. are noct turnal predators of sand beach amphipods, Orchestoidea califomiana (Brandt). I made continuous observations of Thinopinus near the drift patches on which amphipods feed.
2. Thinopinus alternated between active and ambush foraging modes.
3. Thinopinus had greater attack rates and captured smaller amphipods while in active foraging mode.
4. Thinopinus attacked 0.147 amphipods/min, and captured 9.1% of the amphipods attacked. The attack rate on amphipods increased with the proportion of time spent by Thinopinus in active mode.
5. Thinopinus remained longer near types of drift patches which were more attractive to amphipods.     

Slug foraging behaviour: Attraction to food items from a distance

Slug foraging paths recorded using time lapse video techniques under infra-red light were fitted to a correlated random walk model. Analyses were carried out on slugs in arenas where no food was present, as well as those containing commercial molluscicides at two rates of application. Slugs were found to search randomly in empty arenas and those containing commercial metaldehyde baits, but deviated from random search patterns when in the presence of commercial methiocarb bait. Differential attraction from a distance is suggested as a cause of these differences.     

Heritability of corticosterone response and changes in life history traits during selection in the zebra finch

Vertebrates respond to environmental stressors through the neuro-endocrine stress response, which involves the production of glucocorticoids. We have selected independent, duplicate divergent lines of zebra finches for high, low and control corticosterone responses to a mild stressor. This experiment has shown that over the first four generations, the high lines have demonstrated a significant realized heritability of about 20%. However, the low lines have apparently not changed significantly from controls. This asymmetry in response is potentially because of the fact that all birds appear to be showing increased adaptation to the environment in which they are housed, with significant declines in corticosterone response in control lines as well as low lines. Despite the existence of two- to threefold difference in mean corticosterone titre between high and low lines, there were no observed differences in testosterone titre in adult male birds from the different groups. In addition, there were no consistent, significant differences between the lines in any of the life history variables measured--number of eggs laid per clutch, number of clutches or broods produced per pair, number of fledglings produced per breeding attempt, nor in any of egg, nestling and fledgling mortality. These results highlight the fact that the mechanisms that underlie variation in the avian physiological system can be modified to respond to differences between environments through selection. This adds an additional level of flexibility to the avian physiological system, which will allow it to respond to environmental circumstances.     

Pairing context determines condition-dependence of song rate in a monogamous passerine bird

Condition-dependence of male ornaments is thought to provide honest signals on which females can base their sexual choice for genetic quality. Recent studies show that condition-dependence patterns can vary within populations. Although long-term association is thought to promote honest signalling, no study has explored the influence of pairing context on the condition-dependence of male ornaments. In this study, we assessed the influence of natural variation in body condition on song rate in zebra finches (Taeniopygia guttata) in three different situations: during short and long encounters with an unfamiliar female, and within heterosexual mated pairs. We found consistent individual differences in male directed and undirected song rate. Moreover, body condition had a positive effect on song rate in paired males. However, male song rate was not influenced by body condition during short or long encounters with unfamiliar females. Song rate appears to be an unreliable signal of condition to prospective females as even poor-condition birds can cheat and sing at a high rate. By contrast, paired females can reliably use song rate to assess their mate''s body condition, and possibly the genetic quality. We propose that species'' characteristics, such as mating system, should be systematically taken into account to generate relevant hypotheses about the evolution of condition-dependent male ornaments.     

Sex-specific foraging behaviour in a seabird with reversed sexual dimorphism: the red-footed booby

Most hypotheses attempting to explain the evolution of reversed sexual dimorphism (RSD) assume that size-related differences in foraging ability are of prime importance, but the studies on sex-specific differences in foraging behaviour remain scarce. We compare the foraging behaviour of males and females in a seabird species with a RSD by using several miniaturised activity and telemetry loggers. In red-footed boobies males are 5% smaller and 15% lighter than females, but have a longer tail than females. Both sexes spend similar time on the nest while incubating or brooding. When foraging at sea, males and females spend similar time foraging in oceanic waters, forage in similar areas, spend similar proportion of their foraging trip in flight, and feed on similar prey—flying fishes and flying squids—of similar size. However, compared to males, females range farther during incubation (85 km vs. 50 km), and furthermore feed mostly at the extremity of their foraging trip, whereas males actively forage throughout the trip. Males are much more active than females, landing and diving more often. During the study period, males lost mass, whereas females showed no significant changes. These results indicate that males and females of the red-footed boobies differ in several aspects in their foraging behaviour. Although some differences found in the study may be the direct result of the larger size of females, that is, the slightly higher speeds and deeper depths attained by females, others indicate clearly different foraging strategies between the sexes. The smaller size and longer tail of males confer them a higher agility, and could allow them to occupy a foraging niche different from that of females. The higher foraging effort of males related to its different foraging strategy is probably at the origin of the rapid mass loss of males during the breeding period. These results suggest that foraging differences are probably the reason for the differential breeding investment observed in boobies, and are likely to be involved in the evolution and maintenance of RSD.     

Environmental constraints on the foraging behaviour of a dwarf antelope (Madoqua kirkii)

Dik-diks (Madoqua sp.) inhabit semi-arid regions and experience very different conditions of food availability and quality between wet and dry seasons. By comparing the behaviour of dik-diks between these two seasons, we identified environmental constraints affecting their feeding strategies. In both seasons foraging time was limited by high mid day temperatures. In the wet season a high intake rate compensates for the loss in foraging time, but in the dry season water and protein become limiting. To meet minimum daily water requirements in the dry season dik-diks fed on plant species that they avoided during the wet season. Analysis at the plant species level showed higher species selectivity in the wet season than in the dry season. In a multiple regression analysis food species preferences were best explained by relative abundance and water content in the dry season, and by dry matter content in the wet season. In the wet season the daily dry-matter intake of dik-diks in the field was only about 10% higher than the theoretically predicted minimum for a ruminant of this body weight, while protein and water intake were about 3 times as high. This suggests that the most limiting dietary component in the wet season is energy. In the dry season the daily intake of all dietary components is lower than the theoretical minimum required, and also lower than the values suggested by laboratory studies of dik-diks. This dry season deficit is presumably met from body reserves. Dry season water intake was approximately 30% of the intake observed in laboratory studies indicating that dikdiks are even better adapted to arid conditions than suggested by physiological experiments.     

Heritability of Wing Morphology in a Natural Population of Drosophila gouveai

The natural and laboratory heritabilities of a series of parameters related to wing size and shape were estimated in a population of Drosophila gouveai (repleta group) under field and laboratory conditions. A morphometric analysis was done using 17 wing parameters related to wing landmark positions obtained using the method of the best adjustment of an ellipse to the wing edge. Three parameters (thetaA, thetaC and thetaD) showed highly significant heritability in the wild (average 0.61), whereas only wing size (W(SI)) had significant heritability in the laboratory (0.71). The additive genetic variance of most parameters was greater in the wild than in the laboratory. These results showed that some parameters possessed a substantial genetic additive component in their phenotypic variance, and that morphometric parameters of D. gouveai wings are appropriate quantitative markers for assessing morphological differentiation among populations.     

Heavy metal pollution affects dawn singing behaviour in a small passerine bird

Although several studies have suggested that behavioural measures may be more comprehensive than other biomarkers for indicating an organism’s health or welfare, this has rarely been investigated in free-living terrestrial vertebrates. Here we examine the expression of dawn singing behaviour in a free-living small songbird in relation to environmental pollution. We compared the singing behaviour of male great tits Parus major inhabiting an area extremely polluted with heavy metals with that of males inhabiting areas of low(er) pollution (at 4 and 20 km distance from the pollution source). Males at the most polluted site had a significantly smaller repertoire size than males at the two other sites. They also produced a significantly lower total amount of song during the dawn chorus than the males at a distance of 4 km from the pollution source. Our results, although non-experimental and obtained in field conditions, strongly suggest that heavy metal pollution might affect the expression of singing behaviour. Taking into account that previous studies were not able to detect clear, straightforward differences between the health of great tits at the most polluted site and at 4 km distance from the pollution source, our results suggest that the singing behaviour of great tits may be a useful indicator of environmental stress at the population level.     

Short-term cues used by foraging trout in a California stream

Stream salmonids choose foraging locations to maximize the energy benefit of foraging within the constraints of size-mediated dominance hierarchies and predation risk. But, because stream habitats are temporally variable, fish must use a search process to monitor changing habitat conditions as a means of locating potentially-better foraging locations. I explored the cues used by the cutthroat trout, Oncorhynchus clarki clarki, when searching for food at the pool scale by artificially increasing prey availability at different locations by using special feeders and by manipulating pool velocities. Behavior of individually marked fish was monitored from stream bank platforms under unmanipulated control conditions and under seven experimental sets of conditions involving different combinations of feeder location and velocity manipulation. Under natural conditions fish elected to forage in the deepest (>50 cm), fastest (0.10–0.25 m s⁻¹) locations and within 1 m of structure cover, but would readily move to shallower (<30 cm) water away from cover if velocities were manipulated to be highest there. Although fish did not locate feeders unless they were placed in high-velocity areas, when high velocity was provided fish would move into very shallow water (<20 cm) if prey were delivered there. Responses of individual trout to manipulations indicated that water velocity was the main physical cue used by fish to decide where to forage, and that fish could also learn about new food sources by observing conspecifics. Overall, results indicated fish were not “perfect searchers” that could quickly locate new food resources over short time scales, even when the new resources were within a few meters of the fish’s normal foraging location. When given the correct cues, however, fish could detect new food sources and defend them against subordinate fish. Movement of new fish into and out of the study pools during the ten-day observation period was common, consistent with the idea that trout used movement as a means of exploring and learning about habitat conditions at the reach scale.     

Optimal diving behaviour for foraging in relation to body size

Overall, large animals dive longer and deeper than small animals; however, after the difference in body size is taken into account, smaller divers often tend to make relatively longer dives. Neither physiological nor theoretical explanations have been provided for this paradox. This paper develops an optimal foraging diving model to demonstrate the effect of body size on diving behaviour, and discusses optimal diving behaviour in relation to body size. The general features of the results are: (1) smaller divers should rely more heavily on anaerobic respiration, (2) larger divers should not always make longer dives than smaller divers, and (3) an optimal body size exists for each diving depth. These results explain the relatively greater diving ability observed in smaller divers, and suggest that if the vertical distribution of prey in the water column is patchy, there is opportunity for a population of diving animals to occupy habitat niches related to body size.     

Herbivore-induced infochemicals influence foraging behaviour in two intertidal predators

Herbivore-induced defences appear ubiquitous across most biomes and habitats. Yet the direct correlation between induced changes in host plant chemistry and the population dynamics of the herbivore remain untested in many systems. In plant–herbivore interactions in the terrestrial environment, indirect or tritrophic interactions appear a successful way in which changes in the host plant chemistry induced by prior herbivory can impact on herbivore populations via increased success of natural enemies. This set of interactions remains untested in the marine system. Here, we present work from experiments using orthogonal contrasts of plants with different prior treatments (control, mechanical damage or herbivory) and the presence or absence of herbivores on the foraging behaviour of a crab, Carcinus maenas, and a fish, Lipophrys pholis. These experiments were carried out using a novel flow-through flume, i.e. as a choice chamber supplied by turbulent water from independent cue sources. Our results show that in the Ascophyllum nodosum (plant)–Littorina obtusata (herbivore) system infochemicals from induced plants can directly influence predator foraging behaviour. L. pholis was attracted to the presence of a feeding L. obtusata, but was also more attracted to odours from herbivore-induced tissue than odours from mechanically damaged or naïve A. nodosum. C. maenas was more attracted to odours from herbivore-induced tissue compared to naïve tissue, regardless of the presence of L. obtusata. This is the first demonstration of such behavioural consequences of herbivore-induced changes in plants for marine systems.     

Fire induced changes in the foraging behaviour of impala Aepyceros melampus in Lake Mburo National Park, Uganda

The impact of fire on the foraging behaviour of impala (Aepyceros melampus), was studied in the Lake Mburo National Park, Uganda. Two indices of foraging efficiency were used to establish differences in feeding behaviour in burnt and unburnt control areas. These are acceptable grass/browse abundance (AGA/ABA), expressed in terms of feeding time achieved per 50 steps and food ingestion rate (FIR), expressed as the cumulative percentage. Accumulation of above ground grass biomass was also measured in burnt and control areas. After burning, the grazing efficiency remains constant throughout the dry season, while the browsing time achieved per 50 steps decreases drastically. In the control areas AGA decreased with increasing length of the dry season while ABA increased. When expressed in terms of cumulative percentage (FIR), the results show a significant increase of 31% in foraging time spent grazing and a 24% decrease in foraging time spent browsing, in burnt areas. The results confirm the classification of impalas as intermediate or mixed feeders with a distinct preference for grass. Impala foraging patterns after burning are discussed and compared with investigations on the effect of burning on the behaviour of impalas and Buffon's kob (Kobus kob kob) in other areas of Africa.     

Effects of recent experience on foraging in tephritid fruit flies

In field-cage studies, we investigated how the foraging behavior of tephritid fruit flies is modified by experience immediately prior to release on host plants. We observed females of a relatively monophagous species,Rhagoletis mendax (blueberry maggot fly), an oligophagous species,Rhagoletis pomomella (apple maggot fly), and a polyphagous species,Ceratitis capitata (Mediterranean fruit fly). Just prior to release on a host plant, the following kinds of stimuli were supplied: (1) single oviposition in a host fruit, (2) contact with 20% sucrose, (3) contact with a mixture of protein food (bird feces and sucrose), (4) contact with water, and (5) a walk over a host-plant leaf. When flies foraged on host plants without resources, search was most intensive (as measured by number of leaves visited) following a single oviposition in fruit, but residence time generally was the same following exposure to sugar, protein, and fruit stimuli.Rhagoletis mendax andC. capitata females visited the fewest leaves following exposure to water or host leaves, whereasR. pomonella foraged equally intensively following exposure to food stimuli, water, or leaves. On host plants containing resources (fruit and protein food), a single oviposition dramatically increased the number of females of all three species that found fruit compared to females that received experience with food, water or foliar stimuli. We found no significant effect of recent brief experience with any of the stimuli on subsequent attraction to protein food. Overall,C. capitata exhibited a higher propensity to abandon host plants than eitherR. mendax orR. pomonella. We suggest that this may reflect adaptations to differences in distribution of host plants in nature, strategies of dispersal, and host range.