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.     

安氏薮羚的性二型对采食行为的影响(英文)

安氏薮羚(Tragelaphus angasii)是具有性别体二型的动物,其体型大小的差异可能对采食行为和生境选择有显著影响.利用连续行为取样法检测了雌雄个体在生境和植物利用上的差异,记录采食植物种类和采食高度并进行了性别间比较.在每个采食周期末,采用点中心方形样方法(PCQ)对植被进行一次调查,利用判别函数分析鉴别雌雄个体之间在种类和高度上是否存在差异.雌羚在三类生境中花费的采食时间相似,但雄羚表现出对砂地林的偏好.另外,在同一生境内雄羚利用更多的木本种类且采食较高的部位,而雌羚喜欢采食低层草本植物.安氏薮羚雌雄个体在生境内具有食性(采食种类)和空间(生境斑块)上的分离.我们认为这是由性二型引起的性别间营养与能量需求差异和不同繁殖策略造成的,最终使得精食者动物在具有复杂结构空间异质性的生境内表现为空间性别分离     

The influence of food supply on foraging behaviour in a desert spider

We tested the alternative hypotheses that foraging effort will increase (energy maximizer model) or decrease (due to increased costs or risks) when food supply increased, using a Namib desert burrowing spider, Seothyra henscheli (Eresidae), which feeds mainly on ants. The web of S. henscheli has a simple geometrical configuration, comprising a horizontal mat on the sand surface, with a variable number of lobes lined with sticky silk. The sticky silk is renewed daily after being covered by wind-blown sand. In a field experiment, we supplemented the spiders' natural prey with one ant on each day that spiders had active webs and determined the response to an increase in prey. We compared the foraging activity and web geometry of prey-supplemented spiders to non-supplemented controls. We compared the same parameters in fooddeprived and supplemented spiders in captivity. The results support the costs of foraging hypothesis. Supplemented spiders reduced their foraging activity and web dimensions. They moulted at least once and grew rapidly, more than doubling their mass in 6 weeks. By contrast, food-deprived spiders increased foraging effort by enlarging the diameter of the capture web. We suggest that digestive constraints prevented supplemented spiders from fully utilizing the available prey. By reducing foraging activities on the surface, spiders in a prey-rich habitat can reduce the risk of predation. However, early maturation resulting from a higher growth rate provides no advantage to S. henscheli owing to the fact that the timing of mating and dispersal are fixed by climatic factors (wind and temperature). Instead, large female body size will increase fitness by increasing the investiment in young during the period of extended maternal care.     

Temperature constraints on foraging behaviour of male Alpine ibex (Capra ibex) in summer

In arctic and alpine environments, warm summer temperatures may force a reduction in foraging time of large herbivores, whose tolerance for heat is lower than for species adapted to warmer weather. We constructed time budgets for marked ibex (Capra ibex) males over two summers to test whether warm temperatures constrained foraging behaviour and forced altitudinal migrations. As daily temperature and solar radiation increased, feeding activity was reduced at midday and evening, but increased in the early morning, probably to anticipate for an expected reduction in foraging later in the day. With increasing temperature and solar radiation, ibex moved to higher elevations where they spent very little time feeding. Changes in forage quality and availability could not explain altitudinal migration. Temperatures above 15–20°C apparently result in heat discomfort in male Alpine ibex. As temperature and solar radiation increased, older and larger ibex spent less time feeding during daylight and showed a steeper decrease in feeding time than younger and smaller ibex. Larger males may be more sensitive to temperature and solar radiation, or may have more flexibility in allocating time to different activities, given their lower relative energetic requirements. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Host behaviour and its influence on foraging and acceptance by the solitary parasitoid wasp,Venturia canescens (Hym: Ichneumonidae)

Behavioural interactions between the solitary koinobiont parasitoid,Venturia canescens, and two of its hosts,Plodia interpunctella andCorcyra cephalonica, were investigated. The response of both hosts to simulated antennation using a two-haired brush was examined over instars 3 (L3) to 5 (L5). YoungP. interpunctella larvae predominantly adopted escape tactics (writhe, trash) whereas L5P. interpunctella usually froze after the stimulus was applied. L3C. cephalonica larvae were more aggressive (headrear, flick) thanP. interpunctella in response to the application of the stimulus, but olderC. cephalonica responded less aggressively than in earlier instars. AlthoughV. canescens readily jabbed its ovipositor at both hosts after antennation,P. interpunctella was considerably more susceptible to parasitoid attack thanC. cephalonica, irrespective of size in the final (L5) instar.C. cephalonica, the larger, more aggressive host, actively resisted parasitism whereasP. interpunctella responded much more passively after parasitoid contact. Parasitoids examined and jabbed their ovipositors at dead hosts, but this behaviour was not sustained, implying that host movement stimulates parasitoid attack. On patches containingV. canescens, L5C. cephalonica andP. interpunctella, mostP. interpunctella larvae responded by freezing after parasitoid contact.P. interpunctella that froze usually avoided parasitism, whereas larvae that attempted to escape by crawling were pursued with vigour byV. canescens and usually parasitized. Irrespective of behaviour after parasitoid contact,C. cephalonia displayed more aggressive behaviour and had much greater success in warding off parasitoid attack. Host acceptance byV. canescens is clearly affected by the size and species of the host it attacks. The influence of host defensive behaviour is discussed in relation to the evolution of parasitoid counter-defences and oviposition strategies.     

Temporal and spatial pattern of common tern (Sterna hirundo) foraging in the Wadden Sea

In 1989, 313 foraging flights of common terns in the Wadden Sea were radio-tracked. The feeding trips lasted on average 115 min covering about 30 km per flight. Completely tracked flights had a mean radius of 6.3 km. The terns preferred distinct foraging areas in the Wadden Sea. These were visited at site-specific phases of the tidal cycle resulting in a temporal and spatial pattern of foraging, caused by the site-specific and tide-related fluctuations of food availability.Supported by the Deutsche Forschungsgemeinschaft     

Heritability of selectively advantageous foraging behaviour in a small passerine

Summary I measured the heritability of foraging patch choice in a laboratory population of zebra finches (Taeniopygia guttata). Mothers and offspring were tested for their ability to discriminate between four foraging patches which provided four different rates of energy gain. Use of a foraging patch with a high rate of energy gain has been shown to confer a selective advantage on zebra finches in a similar experimental system. In this population of zebra finches there was a large amount of variation in foraging patch choice behaviour both within and among individuals. I determined that foraging patch choice was a phenotypically labile trait with a degree of stereotypy or repeatability, much lower than those typically recorded for morphological traits. The mating behaviour of zebra finches required that heritability be determined from a mother—offspring regression, which showed that narrow sense heritability of foraging patch choice was approximately 0.346. This heritability was significantly different than zero, as was heritability when it was limited by repeatability to 0.246. Foraging patch choice, a behaviour that has a demonstrated fitness consequence, had a heritable component in this laboratory population of zebra finches.     

Foraging activity and behaviour of two goatfish species (Perciformes: Mullidae) at Fernando de Noronha Archipelago, tropical West Atlantic

The goatfishes (Mullidae) include about 50 bottom-foraging fish species. The foraging activity of the yellow goatfish, Mulloidichthys martinicus, and the spotted goatfish, Pseudupeneus maculatus, was studied comparatively at Fernando de Noronha Archipelago, off coast of Northeast Brazil tropical West Atlantic. Pseudupeneus maculatus fed over a larger variety of substrate types, had lower feeding rate, roamed more per given time, spent less time in a feeding event, and displayed a more diverse repertoire of feeding modes than M. martinicus. The differences in the foraging activity and behaviour between the two species possibly minimize a potential resource overlap, as already recorded for other sympatric mullids. Pseudupeneus maculatus had lower feeding rate most likely because it feeds on larger items, and roamed over greater distance per time. Possibly this is because it foraged over a greater variety of substrate distributed over a larger area than that used by M. martinicus. Notwithstanding the overall morphological and behavioural similarity of goatfishes in general, they do differ in their substrate preferences and foraging activity, which indicates that these fishes should not be simply considered generalized bottom foragers.     

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.     

An apparatus and procedure for studying the choice component of foraging in a captive group of chimpanzees (Pan troglodytes)

An apparatus is described which was used to investigate the choice component of foraging in a captive group of chimpanzees maintained in a large, outdoor compound at the Yerkes Regional Primate Research Center Field Station. The utilization of more than one apparatus would allow the investigation of other ecological and psychological concepts in nonhuman primates housed under semi-natural conditions.     

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.     

Physical constraints on the foraging ecology of a predatory snail

We studied the effects of aerial exposure and high summer temperatures on the southern oyster drill ( Stramonita haemastoma ), feeding on the American oyster, Crassostrea virginica . In the laboratory, oyster drill feeding rates and growth were highest at 25 and 30°C, some mortality occurred at 35°C, all snails died at 40 and 45°C, and the 28-day LC 50 was 35.7°C. In a second experiment where both water temperature (25 vs . 33°C) and aerial exposure were varied, only simulated tidal exposure lowered oyster drill feeding and growth rates. In field cage experiments, oyster drills had reduced feeding rates and growth at intertidal sites, but snail growth rates increased in late summer with warmer water temperature. We therefore conclude that aerial exposure, not high temperature, is the major factor limiting oyster drill feeding and growth in intertidal oyster reefs. Field experiments with partial cages also suggested that ambient predation rates were much higher at a subtidal than at a nearby intertidal site. Because southern oyster drills have depressed feeding, growth, and possibly lower fitness in intertidal oyster reefs during the summer, this reduced predation risk may provide a refuge for intertidal oysters.     

Host-plant selection,diet diversity,and optimal foraging in a tropical leafcutting ant

Summary A month-long study was conducted on the comparative foraging behavior of 20 colonies of the leafcutting ant, Atta cephalotes L. in Santa Rosa National Park, Guanacaste Province, Costa Rica. The study was conducted during the middle of the wet season, when trees had mature foliage and the ants were maximally selective among species of potential host plants. The colonies always gathered leaves from more than a single tree species but on average one species constituted almost half the diet with the remaining species being of geometrically decreasing importance. Colonies exhibited greater diversity in their choice of leaves and lower constancy of foraging when the average quality of resource trees was lower, as predicted by elementary optimal foraging theory. Furthermore, the ants were more selective of the species they attacked at greater distances from the nest. However, the ants sometimes did not attack apparently palatable species, and often did not attack nearby individuals of species they were exploiting at greater distances.A classical explanation for why leafcutting ants exploit distant host trees when apparently equally good trees are nearer, is that the ants are pursuing a strategy of conserving resources to avoid long-term overgrazing pressure on nearby trees. We prefer a simpler hypothesis: (1) Trees of exploited species exhibit individual variation in the acceptability of their leaves to the ants. (2) The abundance of a species will generally increase with area and radial distance from the nest, so the probability that at least one tree of the species will be acceptable to the ants also increases with distance. (3) The ants forage using a system of trunk-trails cleared of leaf litter, which significantly reduces their travel time to previously discovered, high-quality resource trees (by a factor of 4- to 10-fold). (4) Foragers are unware of the total pool of resources available to the colony. Therefore once scouts have chanced upon a tree which is acceptable, the colony will concentrate on harvesting from that tree rather than searching for additional sources of leaves distant from the established trail.     

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.     

The effect of prey type and density on the foraging efficiency of juvenile perch, (Perca fluviatilis)

The foraging efficiency of juvenile perch (Perca fluviatilis), feeding on two types of prey, was studied in laboratory experiments. Waterfleas (Daphnia magna) and phantom midge larvae (Chaoborus flavicans) were offered in a range of densities, either separately or combined. Perch fed more efficiently on each prey type separately than when both were mixed. Foraging efficiency decreased with an increase of mixed prey density with both prey types present in equal numbers, but also when the proportion of Chaoborus increased. This could be caused by the existence of different hunting techniques, each of which is fully efficient in the presence of one prey type only. In the presence of two prey types, the predator constantly has to switch from one hunting technique to another.     

Effects of floor-feeding and the presence of a foraging substrate on the behaviour and stress physiological response of individually housed gilts

Both restricted feeding and barren housing have a negative influence on sow welfare. The aim of this study was to test whether sows that have to search for their feed in a substrate on the floor show less stereotyped (and other abnormal) behaviour and have a lower physiological stress response. In three batches, 96 gilts were housed individually in two rooms in 3.1 m² pens with 1.9 m² solid floor. In a 2×2 factorial design either wood shavings (S) or no substrate (NS) were provided on the floor, and 900 g of feed was provided twice daily (06:30 and 15:00 h) either in a trough (T) or on the floor (F). In weeks 8–12, behaviour was scan-sampled once in the periods 07:00–09:00 h (P1), 10:00–12:00 h (P2), and 13:00–15:00 h (P3). Data from the 5 weeks were pooled per animal. Video recordings (24 h) in week 12 or 13 were scan-sampled for ‘standing’. Saliva samples were taken in week 11 at 2 h intervals during 24 h and measured for cortisol. Spontaneously voided morning-urine was sampled in weeks 2, 7 and 12 or 13 for determination of ratios of adrenaline (A) and noradrenaline (NA) with creatinine (CR). Most effects that were found were due to substrate presence. Main findings were that compared with S-animals, NS-animals stood more during the dark period (4.8% versus 3.0%; P<0.05) and showed more (visible) oral behaviour in P1 (56.8% versus 47.6%; P<0.05), P2 (35.8% versus 30.8%; tendency) and P3 (44.1% versus 33.8%; P<0.05). This included more sham chewing in P1 (tendency), P2 and P3, more pen manipulation in P3, and more other oral behaviour (e.g. teeth grinding) in P1 (tendency) and P2. They had higher cortisol levels before feeding (peaks) and in the early evening (24 h average: 1.49 ng/ml versus 1.02 ng/ml; P<0.05) and higher NA/CR ratios in weeks 7 (tendency) and 12 (5.5 ng/mg versus 3.7 ng/mg; P<0.05). No treatment effects were found on A/CR ratios. Substantial interactive effects of feeding method and substrate were only found in floor manipulation. In all three periods NS-T-animals manipulated the floor less than other animals, probably because they had no attraction to the floor and were not rewarded for searching. Results imply that the presence of a substrate on the floor improves welfare, whereas provision of feed in the substrate in order to stimulate foraging behaviour is of less importance.     

The influence of plant architecture on the foraging efficiencies of a suite of ladybird beetles feeding on aphids

By manipulating plant variety and predator species, we investigated the interactions of plant and predator traits in determining predation effectiveness. The predators were all coccinellid adults (Hippodamia convergens, Hippodamia variegata, Coccinella apunctata, and Coccinella septempunctata) and the prey were cabbage aphids (Brevicoryne brassicae). Foraging behavior of the four predators was observed on four crucifers that differed widely in their structures and surface textures (Brassica oleracea caulorapa, Brassica campestris, Brassica juncea crispifolia, and Hirschfeldia incana). Predation rates were significantly influenced by plant variety, a result we attribute to direct effects of plant morphology on predator mobility, falling frequency, and prey accessibility. Predation rates did not vary significantly among the ladybirds, although the four species did exhibit distinct foraging strategies as measured by time spent actively foraging, the rate of encountering aphids, and the fraction of aphids encountered that were consumed. The coccinellids also differed in their propensity for flying away from the plant, and in the frequency with which they fell from the plant. We did not detect any significant interaction effects between plant and predator species, suggesting that the main effects of plant and predator species may overwhelm their interactions in this kind of system. Our results suggest that the level of predation upon herbivorous insects may depend more upon plant architecture than on the particular species of natural enemies present.     

Predation on earwigs: A novel diet specialization within the genusLeptogenys (Formicidae: Ponerinae)

Summary Leptogenys sp. 13 (nearkraepelini) exclusively hunts on earwigs (Gonolabis electa andParalabis sp.). This unusual kind of prey specialization is shown by field data, prey midden investigations, and prey selection experiments. The adaptive value of the species' prey specialization is discussed.     

Magellanic woodpecker ( Campephilus magellanicus) abundance and foraging in Tierra del Fuego, Chile

The Magellanic woodpecker (Campephilus magellanicus) is a vulnerable and poorly studied bird in the sub-antarctic deciduous and evergreen beech (Nothofagus) forests of South America. On Tierra del Fuego island (Chile), we compared Magellanic woodpecker abundance and its foraging habitat in two forest types: pure N. pumilio and mixed forests composed by N. pumilio and N. betuloides, including managed and non managed stands. At a regional scale, abundance of woodpeckers was greater in landscapes including both forest types than in pure N. pumilio landscapes. When both forest types occurred together, woodpecker abundance did not differ between them. The number of trees with foraging signs was correlated with Magellanic woodpecker abundance and was also associated with N. betuloides and snag densities, but was not affected by forest management. Occurrence of pecking on foraging trees was greater in mixed Nothofagus than pure N. pumilio stands. Woodpeckers foraged disproportionately more on larger diameter and more decayed trees. Moreover, trees used for foraging were positively correlated with canopy cover and snag density and were negatively correlated with distance to nearby peatlands and beaver ponds. Direct observation revealed that the flying distance between trees was negatively correlated with proportion of trees with foraging signs. Woodpeckers chose trees that were visited before, suggesting a pattern of tree recognition within foraging territories.Communicated by F. Bairlein     

Homosexual behaviour in wild white-handed gibbons (Hylobates lar)

Homosexual mounting in apes and prosimians is rare. Male-male mounting was observed between an adult male white-handed gibbon and an adolescent male in the same group. The behaviour is discussed in terms of the gibbon group's social structure and the development of the adolescent. It is suggested that this behaviour provides reassurance for the adolescent gibbon.