Prey discrimination by a generalist coccinellid predator: effect of prey age or parasitism?

1. Patterns of prey discrimination by the generalist predatory coccinellid Coleomegilla maculata lengi Timb. (Coleoptera: Coccinellidae) were studied in relation to prey quality, under laboratory conditions. 2. Choice experiments were performed in which second‐ and fourth‐instar coccinellids had the choice between Trichoplusia ni (Lepidoptera: Noctuidae) eggs that were young or old, unparasitised or parasitised by Trichogramma evanescens Westwood (Hymenoptera: Trichogrammatidae). 3. Coleomegilla maculata larvae did not exhibit any preference for parasitised or unparasitised eggs at a similar stage of development but always preferred younger eggs, regardless of whether or not they were parasitised. Furthermore, the percentage of rejection was higher and handling time longer on the less preferred egg type. Pre‐imaginal development time, food intake to reach adulthood, and survival of coccinellid immatures were altered when coccinellid larvae were fed with parasitised and old unparasitised eggs. 4. These results indicate that C. maculata larvae select eggs based on their stage of development regardless of parasitism, and that prey quality of aged prey is lower. 5. Fourth‐instar coccinellid larvae spent less time in patches containing solely parasitised old eggs, and their level of exploitation was greatly reduced, compared with homogeneous patches containing unparasitised young eggs. This suggests that C. maculata larvae respond to variable patch quality by using flexible decision rules that reflect the payoff of the patch.     

Antennal Drumming,Trophallaxis, and Colony Development in the Social Wasp Polistes fuscatus (Hymenoptera: Vespidae)

Parental care is an important component of social behavior in both vertebrates and invertebrates. Social wasps are a useful system for investigating the interplay between behaviors associated with the feeding of larvae by adults and their role in the evolution and maintenance of sociality. Females of the primitively eusocial wasp genus Polistes perform conspicuous vibratory behaviors closely associated with adult–larva feeding interactions. Prior research strongly indicates that these signals are directed toward the larvae, but their function(s) remain unclear. Existing hypotheses on the function(s) have posited releaser effects on larvae, either stimulating or inhibiting release of larval saliva, a nutrient‐rich glandular secretion attractive to adults. Polistes fuscatus queens perform antennal drumming (AD), a behavior in which they rapidly beat their antennae synchronously on the rims of the nest cells during the feeding of larvae. We used radiolabeled prey to show that adults extract juice from the prey, which they subsequently regurgitate to larvae immediately following each AD burst. We also show that no saliva is imbibed by the queen during the contact. These results are consistent with the inhibition hypothesis on the function of AD, but not the stimulation hypothesis. We further demonstrate that AD is first performed on nests when the oldest larvae reach the third instar, and that the third instar is the first to produce measurable volumes of larval saliva. Removal of third‐, fourth‐, and/or fifth‐instar larvae from single‐foundress, pre‐pupal‐stage colonies did not cause a reduction in the queen’s AD rates compared with controls, suggesting that later‐stage larvae do not maintain AD behavior via an immediate releaser effect. We propose instead that third‐instar larvae, possibly via chemical components of the salivary secretion itself, modulate the physiology of queens so as to indirectly cause the onset and maintenance of AD behavior.     

Mothers vigilantly guard nests after partial brood loss: a cue of nest predation risk in a paper wasp

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Ontogenetic shifts in the functional response and interference interactions of Rhyacophila dorsalis larvae (Trichoptera)

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Therefore, it is important to establish if the functional response and interference interactions differ between life‐stages. These hypotheses were tested by (i) comparing the functional response of second, third, fourth and fifth larval instars of Rhyacophila dorsalis, using three stream tanks with one Rhyacophila larva per tank and one of 10 prey densities between 20 and 200 larvae of Chironomus sp.; (ii) using other experiments to assess interference within instars (two to five larvae of the same instar per tank), and between pairs of different instars (one, two or three larvae per instar; total predator densities of two, four or six larvae per tank). 2. The first hypothesis was supported. The number of prey eaten by each instar increased with prey density, the relationship being described by a type II model. The curvilinear response was stronger for fourth and fifth instars than for second and third instars. Mean handling time did not change significantly with prey density, and increased with decreasing instar number from 169 s for fifth instars to 200 s for second instars. Attack rate decreased progressively with decreasing instar number. Handling time varied considerably for each predator–prey encounter, but was normally distributed for each predator instar. Variations in attack rate and handling time were related to differences in activity between instars, fourth and fifth instars being more active and aggressive than second and third instars, and having a higher food intake. 3. The second hypothesis was partially supported. In the interference experiments between larvae of the same instar or different instars, mean handling time did not change significantly with increasing predator density, and attack rate did not change for second and third instars but decreased curvilinearly for fourth and fifth instars. Interference between some instars could not be studied because insufficient second instars were available at the same time as fourth and fifth instars, and most third instars were eaten by fourth and fifth instars in the experiments. Prey capture always decreased with decreasing attack rate. Therefore, interference reduced prey consumption in fourth and fifth instars, but not in second and third instars. The varying feeding responses of different instars should be taken into account when assessing their role in predator–prey relationships in the field.     

Prey stage preference and feeding behaviour of Oligota kashmirica benefica (Coleoptera: Staphylinidae), an insect predator of the spider mite Tetranychus urticae (Acari: Tetranychidae)

We studied the prey stage preference and feeding behaviour of the first to third instar larvae and adult females of Oligota kashmirica benefica Naomi (Coleoptera: Staphylinidae), a predator of the spider mite Tetranychus urticae Koch (red form) (Acari: Tetranychidae), on leaves of the kudzu vine (Pueraria lobata (Wild.) Ohwi (Leguminosae)) under laboratory conditions. The number of mites eaten increased with the growth of predator larvae. Third instar larvae preyed on all stages of spider mite, whereas first instar larvae preyed mainly on immobile stages (eggs and quiescent stages). The predator larvae showed two types of foraging behaviour (active searching and ambush behaviour) when targeting the mobile stages (larval, nymph and adult stages of prey). Although no difference was found in the number of prey consumed by adult females and third instar larvae of the predator, the adult females mainly attacked and consumed the immobile stages.     

Prey stage preference and feeding behaviour ofOligota kashmirica benefica (Coleoptera: Staphylinidae), an insect predator of the spider miteTetranychus urticae (Acari: Tetranychidae)

We studied the prey stage preference and feeding behaviour of the first to third instar larvae and adult females ofOligota kashmirica benefica Naomi (Coleoptera: Staphylinidae), a predator of the spider miteTetranychus urticae Koch (red form) (Acari: Tetranychidae), on leaves of the kudzu vine (Pueraria lobata (Wild.) Ohwi (Leguminosae)) under laboratory conditions. The number of mites eaten increased with the growth of predator larvae. Third instar larvae preyed on all stages of spider mite, whereas first instar larvae preyed mainly on immobile stages (eggs and quiescent stages). The predator larvae showed two types of foraging behaviour (active searching and ambush behaviour) when targeting the mobile stages (larval nymph and adult stages of prey). Although no difference was found in the number of prey consumed by adult females and third instar larvae of the predator, the adult females mainly attacked and consumed the immobile stages.     

Predation on Mosquito Larvae by Mesocyclops thermocyclopoides (Copepoda: Cyclopoida) in the Presence of Alternate Prey

The cyclopoid copepod Mesocyclops thermocyclopoides, a dominant invertebrate predator in many shallow ponds and temporary water bodies in northern India, feeds on cladocerans, rotifers, ciliates and when present, on mosquito larvae also. We studied in the laboratory the prey consumption rates of the copepod on first and fourth instar larvae of two species of mosquito (Anopheles stephensi and Culex quinquefasciatus) in relation to their density. We also studied its prey selectivity with mosquito larvae in the presence of an alternate prey (the cladocerans‐either Moina macrocopa or Ceriodaphnia cornuta) in different proportions. With either mosquito species, the copepod actively selected Instar‐I larvae, avoiding the Instar‐IV larvae, and with either instar, selected Anopheles stephensi over Culex quinquefasciatus. When prey choice included the cladoceran as an alternate prey, the copepod selected the cladoceran only when the other prey was Instar‐IV mosquito larvae. Our results point to the potential and promise of M. thermocyclopoides as a biological agent for controlling larval populations of vectorially important mosquito species.     

Prey switching in Rhyacophila dorsalis (Trichoptera) alters with larval instar

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Previous studies have shown that the diel activity pattern and functional response differed between larval instars of the carnivorous caddis, Rhyacophila dorsalis. The present study examines switching by larvae of R. dorsalis presented with different proportions of two prey types; either small (length 2–4 mm) and large (5–8 mm) Chironomus larvae for second, third, fourth and fifth instars of R. dorsalis; or Baetis rhodani (9–12 mm) and large Chironomus larvae for fourth and fifth instars. Experiments were performed in stream tanks with one Rhyacophila larva per tank and 200 prey arranged in nine different combinations of the two prey types (20 : 180, 40 : 160, 60 : 140, 80 : 120, 100 : 100, 120 : 80, 140 : 60, 160 : 40 and 180 : 20). Prey were replaced as they were eaten. A model predicted the functional response in the absence of switching and provided a null hypothesis against which any tendency to switch could be tested. 2. There was no prey switching in the second and third instars, with both instars always showing a preference for small over large Chironomus larvae. Prey switching occurred in the fourth and fifth instars. As the relative abundance of one prey type increased in relation to the alternative, the proportion eaten of the former prey changed from less to more than expected from its availability, the relationship being described by an S‐shaped curve. In the experiments with small and large Chironomus, the two instars switched to large larvae when their percentage of the total available prey exceeded 29% and 37% for fourth and fifth instars, respectively. In the experiments with Baetis and large Chironomus, both instars switched to Baetis larvae when their percentage of the total available prey exceeded 36%. 3. Non‐switching in second and third instars was related to their feeding strategies, both instars preferring smaller prey items. When the fourth and fifth instars foraged actively at night, they preferred larger over small Chironomus larvae, but when they behaved as ambush predators at dusk, they captured the more active Baetis larvae in preference to the more sedentary large Chironomus larvae and only switched to the latter when they were >64% of the available prey.     

Prey Selection by the Lady Beetle Harmonia axyridis: The Influence of Prey Mobility and Prey Species

The influence of prey mobility and species on prey selection by the coccinellid Harmonia axyridis Pallas was determined under laboratory conditions for two prey species, Hyaliodes vitripennis (Say) and Tetranychus urticae Koch. Prey selection was influenced by prey mobility. In the presence of active prey, the coccinellid selected T. urticae while in presence of immobilized prey, H. vitripennis was preferred. Harmonia axyridis searching time was longer in the presence of active H. vitripennis than in the presence of active T. urticae. Moreover, the coccinellid capture rate was lower for active H. vitripennis caused by effective defensive mechanisms. Prey suitability was affected by prey mobility and species. Immobilized H. vitripennis were the most profitable prey, i.e. induced a shorter developmental time and no mortality. However, active H. vitripennis were not a suitable food source for H. axyridis. Our results suggested that three factors are involved in prey selection by H. axyridis: (i) prey mobility; (ii) prey defensive mechanisms; and (iii) prey species.     

Larval oophagy in the ant <Emphasis Type="Italic">Amblyopone silvestrii</Emphasis> (Hymenoptera,Formicidae)

Summary Larvae pass through five instars in the temperate, subterranean ponerine ant, Amblyopone silvestrii. Field colonies displayed a large decrease in the number of eggs during mid-summer, despite the fact that queens maintained fully developed ovaries with mature eggs. Observations of laboratory colonies indicate that cannibalism by 1st and 2nd instar larvae caused this decrease in egg number. These instars consumed a total of 66–75% of eggs in the nest, with each larva consuming 2–3 eggs before molting to the 3rd instar. At that time the larvae began to feed on arthropods. The preferred prey of A. silvestrii consists of entire centipedes; the large size of these prey items relative to the size of early instar larvae makes it difficult for the larvae to feed on these prey. Additionally, workers of A. silvestrii do not engage in oral trophallaxis. Consequently, oophagy is a plausible method to feed these very small larvae.     

Contrasting diel activity and feeding patterns of four instars of Rhyacophila dorsalis (Trichoptera)

1. Ontogenetic shifts in prey choice and predator behaviour can affect food‐web structure. Therefore, it is important to establish if the diet and feeding activity differ between life‐stages of the same species. This hypothesis was tested for second, third, fourth and fifth larval instars of Rhyacophila dorsalis by comparing their diel activity and feeding patterns. Second to fifth instars collected from two streams were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks. Food was provided in excess; being organisms living in bryophytes on top of a large stone in each tank, augmented by different‐sized larvae of Ephemeroptera, Simuliidae and Chironomidae. As few first instars for gut analyses were found in the field, the diet of first instars reared in the laboratory was also studied. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 50 larvae per instar for each day or night sample). First and second instars fed on the smaller food items with no significant day‐night differences in diet. Gut contents indicated a progressive trend from feeding chiefly at night in third instars to almost exclusively at night in fifth instars. Fourth and fifth instars fed on the larger food items, whilst the diet of the third instar larvae overlapped with that of both the earlier and later instars. 3. Diel activity patterns of single larvae differed between instars but not within each instar (n = 20 larvae per instar). Second instars were active throughout the 24 h, with peaks at dusk, around midnight, dawn and around midday. A similar pattern was shown by third instars but the peak of activity at midday was less than the other three peaks. Prey were captured only during these peaks for both instars. Fourth and fifth instars were most active, and fed only, at night. They used an ambush strategy to capture more active prey at dusk and dawn (e.g. Baetis, Gammarus), and a searching strategy to capture more sedentary prey during the night (e.g. chironomids, simuliids). These experiments provided support for the hypothesis under test. If competition and/or interference occur between instars, then it could be reduced between earlier and later instars because of differences in their diet and diel pattern of feeding activity.     

Early collapse of Vespa simillima (Hymenoptera, Vespidae) colonies in central Japan

During the past 15 years in the Ina Valley, Japan, mature fifth instar larvae of the yellow hornet, Vespa simillima, have been occasionally ejected from colonies during September. During 2005, this unusual behavior was particularly widespread, and collection of several V. simillima colonies confirmed that very few fifth instar larvae were present. When compared with an average colony, constructed from 41 colonies collected 20 years previously, colonies in 2005 had 80% fewer fifth instar larvae, despite queen egg‐laying rates remaining similar. It may be that the amount of food provided to the larvae by workers is less than the amount of larval secretions received by workers, which is causing the larvae to become emaciated and preventing them from pupating. This phenomenon normally occurs naturally at the end of the colony cycle in November, when prey abundance decreases and larval secretions or other sources of carbohydrates, for example from ripe fruit, tree sap, or aphid secretions, are needed by sexuals to build up fat reserves. However, another possibility is that the unexplained appearance of this phenomenon in September, which is causing colonies to collapse without producing any or very few sexuals, is due to foragers feeding the larvae prey that are contaminated with insect growth regulators, via pesticides, which are known to prevent successful pupation.     

Differential vulnerability determines the diet of a slow-moving predatory stream insect

1. We investigated the diet and prey electivity of Rhyacophila obliterata, a slow‐moving invertebrate predator capable of hunting in high‐flow microhabitats, and quantified the components of the predation sequence of fifth‐instar larvae foraging on mobile (Baetis mayflies, Amphinemura stoneflies) versus semi‐sessile (larval blackflies) prey. 2. In the field, fifth‐instar Rhyacophila consistently took more larval blackflies than more mobile prey. In behavioural trials, the number of attacks by Rhyacophila differed significantly between prey types, mobile prey being attacked more often than blackflies. Capture success, by contrast, was highest for blackflies, whereas Amphinemura and Baetis were rarely captured. In mixed‐prey feeding trials, Rhyacophila showed strong preference for blackflies and equally strong avoidance of Amphinemura and Baetis. 3. For mobile prey, the risk of being captured by this sluggish predator is very low, so they can afford to be in close contact with it. Rhyacophila was almost unable to capture any other prey but blackflies, resulting in strong passive selection for blackflies. 4. Therefore, the diet of fifth‐instar Rhyacophila can be predicted from laboratory observations and prey behaviour is the major determinant of the diet of this invertebrate predator.     

Resource Management in the Orb-Weaving Spider Zygiella x-notata: I. Influence of Spider Behaviour and Available Prey

Management of resource exploitation by the orb-weaving spider Zygiella x-notata was investigated in relation to available prey. Prey capture is studied according to prey type (cricket or fly), and the spider's behaviour (waiting without prey or consuming other prey), at the moment when a prey item was placed onto the web. Characteristics of the spider's exploitation of the previous item (feeding time, and quantities ingested) were analysed according to the type and quantity of prey presented, (1 or 2 items at 1-h intervals). Capture time did not vary in relation to prey type, nor spider activity. However, duration and quantities of certain behavioural components (biting and wrapping) did vary according to prey type and the spider's behaviour, at the moment of presentation of prey. Whatever the type of prey received, if the first item consumed was a cricket, the spiders reduced time spent feeding on this prey without any decrease in the quantities ingested.     

Development of Orius vicinus (Ribaut) (Heteroptera: Anthocoridae) on Different Prey

The survival, development and adult size of Orius vicinus were studied on five prey species, the mites Aculus schlechtendali, Panonychus ulmi and Tetranychus urticae, and larvae of the insects Dasineura mali and Thrips obscuratus. Survival was high (74-100%) and O. vicinus was able to complete development on all prey species in an average of 22-26 days at 20 C. The most rapid development occurred on T. obscuratus, T. urticae and P. ulmi. Adult size was estimated from the length of the fifth instar cast skin, which averaged 2.18 to 2.48 mm on different prey. The largest bugs were obtained from feeding on T. obscuratus or D. mali. O. vicinus, which were fed A. schlechtendali, were the slowest to develop and produced the smallest adults. The results are discussed in relation to prey availability and observed predation in the field, and are compared with previous feeding studies.     

Resource Management in the Orb-Weaving Spider Zygiella x-notata II. Ability to Manage a Set of Various Prey

Management of a set of various prey received ‘simultaneously’ by the orb-weaving spider Zygiella x-notata is investigated in relation to prey species, weight, order of arrival in the web, and factors concerning the spider's state of nourishment. When the spiders encountered several different prey ‘simultaneously’, they subdued all of them, took one of them to begin consumption, and stored the others in the web. The first item for consumption was chosen according to the characteristics of the available prey. Factors concerning previous feeding influenced the spiders's choice of the first item to be ingested. Exploitation of the first item, estimated by the quantity of food extracted and the time necessary for this extraction, depended on the type and the quantity of prey at the spider's disposal. When the spiders received several prey, they spent less time feeding on the first cricket compared with spiders receiving only one cricket, while no difference was found when the first prey item exploited was a fly.     

Nest attendants in parakeet auklets

Breeding behaviors in a captive population of parakeet auklets (Cyclorrhynchus psittacula) were observed to determine whether a third bird present at the entrances to nest tunnels was a helper or competitor. Comparisons were made between focal pairs of nesting birds, two with and two without attendants. Pairs with attendants spent more time feeding and more time bathing and had less time with intruders (other than the third bird) at their tunnels. Nest attendants also helped in nest defense and incubation (taking some of the burden off the focal pair). Relatedness is unknown; therefore, kin selection cannot be assessed. Because all nest tunnels were occupied, however, limited breeding resources in the captive environment may make helping an adaptive alternative for mature birds. Zoo Biol 20:271–277, 2001. © 2001 Wiley‐Liss, Inc.     

Combined effects of allelochemicals, prey availability, and supplemental plant material on growth of a generalist insect predator

We examined the effects of the presence of plant allelochemicals in prey diet, prey availability and supplemental plant material on the growth of the generalist predator Podisus maculiventris (Hemiptera: Pentatomidae). We tested two different nymphal stages of this predator. Third to fourth instar nymphs and fifth instar nymphs were fed a diet of prey (Manduca sexta larvae, Lepidoptera: Sphingidae) without allelochemicals in their diet or prey fed maximal levels of allelochemicals (tomatine, rutin and chlorogenic acid) found in their host plant (Lycopersicon esculentum). The nymphs were fed prey ad libitum, once every three days, or once every five days. They were given either no supplemental plant material or a 2 cm slice of green bean pod (Phaseolus vulgaris). We also conducted another experiment with fifth instar nymphs using the same conditions, except that mean levels of allelochemicals found in the host plant were fed to prey instead of maximal levels and the prey were provided either once a day or once every five days. For all experiments, prey scarcity depressed developmental rate, weight gain and relative growth rate. Overall, there was no negative effect of allelochemicals in the diet of the prey on these variables when predators were supplied with an excess of prey, but allelochemicals in the prey diet negatively affected these predators when prey were scarce. The addition of plant material to the diet of third to fourth instar nymphs did not have any effect on developmental rate, final dry weight, or relative growth rate. However, for fifth instar nymphs, the addition of plant material negatively affected these variables. Thus, the addition of plant material to the diet of the nymphs did not alleviate the negative effects of prey scarcity or allelochemicals in prey diet.     

On “intercolonial” cannibalism in Japanese paper wasps,Polistes chinensis antennalis Pérez andP. Jadwigae dalla torre (hymenoptera: Vespidae)

Summary Foundresses of two species of Japanese paper wasps,Polistes chinensis antennalis andP. jadwigae, attacked other colonies of the same species. A foundress ofP. chinensis antennalis visited two nests of the same species, and ate larvae from them, while two foundresses ofP. jadwigae each visited a nest of the same species, eating larvae and pupae even when the foundress of the attacked nest was on her nest. In addition, a foundress ofP. jadwigae distributed flesh balls thus obtained among their larvae. Discussion was made on the adaptive significance of the inter-colonial cannibalism. It was considered that, at first, it increases the foraging efficiency and secondly it plays a role in regulating population density.     

Time constraint on food choice in provisioning blue tits, Parus caeruleus: the relationship between feeding rate and prey size

Previous work on food-provisioning behaviour in blue tits suggested that the parents could gather larger prey items only by making longer foraging excursions, for example, by being more selective or by reaching more distant (and less exploited) feeding sites. Here, I show that within-nest, within-day variation in size of prey delivered by the parent could be explained by the time since its last visit. In unmanipulated conditions, size of larvae tended to increase with the time spent away from the nest. A significant positive relationship was more likely at high provisioning rates, suggesting that periods of intense feeding limited the size of prey delivered to the brood. To assess the effect of less intense feeding on prey size, I experimentally increased food availability to the tits. The parents could decide whether to eat the extra food or feed it to the nestlings. In both cases, food supplementation could result in longer time lags between natural feedings. Food-supplemented parents consumed the extra food and fed it to their nestlings, made longer foraging trips and delivered larger natural larvae than controls. In this group, size of larvae was more constant during the observation period and was independent of the time since the parent's last visit. This suggests that, below some value of visit rate, prey size is no longer limited by the duration of the foraging trip. The results support the view that tits continually vary visit rate and prey size. There is some evidence that these adjustments are made by changing food selectivity in response to changes in the state of the brood and of the parents.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .