The importance of location and visual cues during foraging in the German wasp (Vespula germanica F.) (Hymenoptera: Vespidae)

Abstract

The way in which foraging wasps use cues for prey location and choice appears to depend on both the context and on the type of prey. Vespula germanica is an opportunistic, generalist prey forager, and individual wasp foragers often return to hunt at sites of previous hunting success. In this paper, we studied which cues are used by this wasp when relocating a food source. Particularly we analysed the response to a displaced visual cue versus a foraging location at which either honey or cat food had been previously presented. We conclude that location is used over a displaced visual cue for directing wasp hovering, although the landing response is directed differently according to bait type. When wasps are exploiting cat food, location also elicits landing, but if they are exploiting honey, a displaced visual cue elicits landing more frequently than location.     

A latitudinal gradient in intensity of applying ant-repellent substance to the nest petiole in paper wasps (Hymenoptera: Vespidae)

Summary Observations on pre-emergence, single-foundress colonies of Japanese paper wasps (Polistes) revealed that there was a latitudinal gradient in intensity of application of an ant-repellent substance (secreted by the metasomal sternum VI glands) to the nest petiole. Thus the lower the latitude, the more frequently a foundress rubbed ant repellent onto the nest petiole. Estimation of potential ant predation on wasp brood using bait traps showed that there was a positive correlation between the frequency of rubbing and potential predation pressure from ants which are guided predominantly by substrate cues for foraging. There was also a latitudinal gradient in the degree of temporal association of rubbing behavior with foraging: the lower the latitude, the more closely foundress departure from the nest was associated with rubbing. Enlargement of the nest petiole by applying oral secretion potentially obliterated previous coats of ant repellent; however, this behavior was not always followed by rubbing behavior. The ant-repellent chemical barrier around the nest petiole may have evolved in tropical regions of the world as a defense against ant predation on wasp brood. I argue that as ant predation pressure diminishes towards the cooler regions, so does selection maintaining the behavioral sequence where foundress departure from the nest is preceded by rubbing behavior.     

海切叶蜂的筑巢和访花行为

【目的】研究毛乌素沙地重要野生传粉昆虫海切叶蜂Megachile maritima的筑巢和访花行为对保护其栖息环境具有重要意义。【方法】采用目测及拍照等方法对海切叶蜂的整个筑巢过程进行了连续观测;以2 m×2 m 样方的方式观测海切叶蜂的访花频率、单花停留时间及日活动规律等访花行为,其中日活动规律每天连续观测,共观测7天。【结果】海切叶蜂在沙土中筑巢,每巢只有一个巢室,其筑巢过程为：寻找合适的筑巢地点,挖巢,构建巢室,采集蜂粮,产卵,封住巢室,筑完一个巢。它连续筑完一个巢大约需要9 h。海切叶蜂构建一个巢室需要切取26～29片叶子,为每巢室采集蜂粮11～12次,每巢室内产卵1粒;在塔落岩黄芪和细叶益母草上的平均访花频率分别为（13.23±6.49）朵/min和（16.72±4.84）朵/min,平均单花停留时间分别为（3.08±2.48）s和（2.49±1.31）s。晴天,海切叶蜂在12:00～14:00期间活动较活跃。【结论】海切叶蜂不同个体之间的筑巢行为相似。该蜂在塔落岩黄芪和细叶益母草上的访花过程、访花频率及单花停留时间具有显著的差异。     

Ground nesting in the paper wasp <Emphasis Type="Italic">Polistes aurifer</Emphasis> (Hymenoptera,Vespidae)

Summary Polistes paper wasp species vary in their nest site selection, with some nesting in relatively exposed areas and others in protected cavities. Locating the nest sites of cavity dwelling species in natural habitat can prove difficult, and most behavioral studies on these species are therefore conducted using human-built structures. Since Hungerford and Williams (1912) and Rau (1929) noted the location of several P. fuscatus (Fabricius) nests in rodent burrows, there have been no published accounts of ground nesting Polistes species. I report the occurrence of a large, dense cluster of P. aurifer (Saussure) nests located within cracks in the dried soil of the Santa Monica Mountains of southern California, USA.Received 8 April 2003; revised 10 October 2003; accepted 31 October 2003.     

Sociogenetic structure of Polistes (Aphanilopterus) versicolor Olivier, 1791 colonies (Hymenoptera, Vespidae, Polistini)

The observation of two distinct, well-defined oviposition areas in nests of the primitively eusocial wasp Polistes versicolor suggests the presence of multiple egg-layers and territorial behaviors. Electrophoretic analysis of enzyme loci in pupae from 35 colonies revealed an average observed heterozygosity of 0.10 and the existence of private polymorphisms, thereby indicating a low dispersion in this species. No evidence of diploid males was found. Phenotypic segregation analysis revealed the presence of more than one egg-laying female in 15 out of 35 colonies, as well as spatially preferential oviposition in 2 out of 13 nests, with distinct oviposition areas. Genetic relatedness estimates for brood were lower than expected for haplodiploid species under monogynous conditions (r = 0.75 for female broods and r = 0.5 for male) in 4 of those 13 nests, thereby inferring complex sociogenetic structuring in Polistes versicolor colonies.     

长木蜂的筑巢和采粉贮粮行为

【目的】通过研究长木蜂Xylocopa tranquebarorum筑巢和贮粮行为, 为进一步查明独栖性蜂类行为特点、开发新的蜂类资源提供依据。【方法】采用目测和拍照等方法对长木蜂的整个筑巢过程进行了连续观察, 用游标卡尺对巢口大小进行测量, 采用室内解剖巢室对长木蜂贮蜂粮的大小和数量进行观测。【结果】长木蜂主要在竹子上筑巢, 偶尔也发现在芦苇上筑巢。最喜欢选择竹节直径1.2～2.5 cm的孝顺竹Bambusa multiplex和刚竹属 Phyllostachys的竹种上筑巢。其筑巢过程为：雌蜂寻找合适的筑巢地点, 咬巢口, 清理巢室, 采集花粉蜜制作蜂粮, 在蜂粮上产卵, 制作巢室隔板。筑巢地点主要位于离旧巢1 m以内的位置。雌蜂啃咬巢口平均用时（292±29）min, 制作一块蜂粮需采集粉蜜22～40次, 采集粉蜜平均用时（17.31±0.52）min/次, 携粉蜜回巢滞留时间平均为（16.45±1.08）min/次;巢中卸落粉蜜平均用时为（15.29±1.03）min/次, 一生贮蜂粮平均6块左右;蜂粮近长方形, 长12～18 mm, 宽6～10 mm, 平均重量（0.7140±0.0269）g。【结论】长木蜂雌蜂不同个体之间筑巢行为相似, 而采集粉蜜的次数和贮蜂粮所用时间均有显著差异。     

Temporal relationships of rubbing behavior with foraging and petiole enlargement inParapolybia indica (Hymenoptera: Vespidae)

Summary The frequency of application of ant repellent secreted from the metasomal sternum VI gland to the nest petiole (=rubbing behavior) inParapolybia indica colonies decreased significantly from the preemergence, single-foundress stage to the post-emergence stage. In the pre-emergence stage, rubbing occurred in close temporal association with the departure from the nest, which may maximize the antrepelling effect of the rubbing substance during the period when the nest is not attended by any wasps. On the other hand, only a small proportion of the departures was associated with rubbing in the post-emergence stage.Unlike in other independent-founding polistine wasps, rubbing inP. indica was not associated with petiole enlargement. Among independent-founding polistines,P. indica is unusual in that females enlarge the nest petiole with a large amount of plant fibers instead of strict use of adult saliva, and in that the petiole surface is rough, and may be much more absorbent than that of the secretion petiole. On the other hand, in the species showing a temporal association between petiole enlargement and rubbing, females typically rub immediately after petiole licking (application of oral secretion). This fact, and the lack of such a temporal association inP. indica, may suggest a possibility that the mixture of oral secretion and rubbing substance on the smooth surface of the secretion petiole makes the chemical barrier last longer.     

Hole repair and the influence of learning on nest repair in the paper wasp,Polistes fuscatus (Hymenoptera: Vespidae)

Two methods of hole repair used by the paper wasp, Polistes fuscatus,were examined and compared, and the hypothesis that nest repair behavior may improve with experience was tested. Similarly sized holes were repeatedly made in the sides of nests and the repair behavior of the nest residents was recorded. The distance of the hole from the cell mouth determined which repair method was used. In addition, both the amount of time for hole repair completion and the number of pulp loads used in making the repair decreased significantly with experience.     

Distributed cognition and social brains: reductions in mushroom body investment accompanied the origins of sociality in wasps (Hymenoptera: Vespidae)

The social brain hypothesis assumes the evolution of social behaviour changes animals'' ecological environments, and predicts evolutionary shifts in social structure will be associated with changes in brain investment. Most social brain models to date assume social behaviour imposes additional cognitive challenges to animals, favouring the evolution of increased brain investment. Here, we present a modification of social brain models, which we term the distributed cognition hypothesis. Distributed cognition models assume group members can rely on social communication instead of individual cognition; these models predict reduced brain investment in social species. To test this hypothesis, we compared brain investment among 29 species of wasps (Vespidae family), including solitary species and social species with a wide range of social attributes (i.e. differences in colony size, mode of colony founding and degree of queen/worker caste differentiation). We compared species means of relative size of mushroom body (MB) calyces and the antennal to optic lobe ratio, as measures of brain investment in central processing and peripheral sensory processing, respectively. In support of distributed cognition predictions, and in contrast to patterns seen among vertebrates, MB investment decreased from solitary to social species. Among social species, differences in colony founding, colony size and caste differentiation were not associated with brain investment differences. Peripheral lobe investment did not covary with social structure. These patterns suggest the strongest changes in brain investment—a reduction in central processing brain regions—accompanied the evolutionary origins of eusociality in Vespidae.     

Evidence of alarm pheromones in the venom of Polistes dominulus workers (Hymenoptera: Vespidae)

Abstract.  The active and coordinating capacity of defending the nest is a key feature of social insects. The present study investigates the presence of alarm pheromones in the venom of workers of the social wasp, Polistes dominulus . Laboratory experiments were performed with caged colonies of P. dominulus using a wind tunnel apparatus to test the behavioural response of workers to venom released by other workers and to venom extracts. Contrary to that previously reported for European paper wasps, the present results show that the venom is the source of alarm pheromones. Field experiments combining a visual (black target) and a chemical stimulus (venom extract) were performed to test the effect of the venom on the reaction of colonies. Wasps leave the nest, land on the visual target and attack the target significantly more once exposed to venom extract plus target than to solvent plus target. This work shows that the venom of P. dominulus workers elicits an alarm response, reduces the threshold for attack and acts as an attractant on targets. These results using P. dominulus indicate that, in both American and European species, colony defence is based on the same features, suggesting that chemical alarm is a widespread trait in the genus Polistes .     

Venom spraying inParachartergus colobopterus: A novel defensive behavior in a social wasp (Hymenoptera: Vespidae)

Colonies ofParachartergus colobopterus do not defend against vertebrates by attacking and stinging. Instead, defending workers bend the gaster forward and spray a fine mist of venom in the direction of nearby moving objects. Although venom spraying has been reported forP. fraternus, a species that does sting, we found that this occurred only during sting attempts. We conclude that defensive behavior inP. colobopterus is unique among wasps in that (1) venom is sprayed at intruders by workers standing on the nest and (2) the spray is an atomized mist rather than a stream. We suggest that nest crypticity restricts vertebrate predators on this wasp to small, insect gleaning birds, against which a spray of venom in the eyes, mouth, and nasal passages is more effective than is a stinging defense.     

斑痣悬茧蜂对不同寄主密度斑块的选择和最优搜寻行为

在圆柱形透明有机玻璃罩内,观察了斑痣悬茧蜂对不同寄主密度斑块的选择和滞留时间,并用Cox比例风险模型分析了寄主密度、产卵次数和搜寻特征对斑痣悬茧蜂在斑块上的驻留时间及其影响因素.结果表明:斑痣悬茧蜂对高密度寄主斑块的初次选择次数、访问次数均高于低密度寄主斑块,导致在高密度斑块上具有更高的寄生率;斑痣悬茧蜂在各斑块的驻留时间随寄主密度的增加而增加.Cox模型分析结果显示,斑块内寄主密度和产卵数均对其离开斑块的倾向起消极作用,即随寄主密度和产卵数的增加而倾向于留在斑块上;而寄主密度与产卵数的互作反而对寄生蜂离开斑块倾向具有促进作用,这两种作用可使寄生蜂利用寄主斑块获得最大适合度.     

Long‐term spatial memory in Vespula germanica social wasps: the influence of past experience on foraging behavior

Social insects exhibit complex learning and memory mechanisms while foraging. Vespula germanica (Fab.) (Hymenoptera: Vespidae) is an invasive social wasp that frequently forages on undepleted food sources, making several flights between the resource and the nest. Previous studies have shown that during this relocating behavior, wasps learn to associate food with a certain site, and can recall this association 1 h later. In this work, we evaluated whether this wasp species is capable of retrieving an established association after 24 h. For this purpose, we trained free flying individuals to collect proteinaceous food from an experimental plate (feeder) located in an experimental array. A total of 150 individuals were allowed 2, 4, or 8 visits. After the training phase, the array was removed and set up again 24 h later, but this time a second baited plate was placed opposite to the first. After 24 h we recorded the rate of wasps that returned to the experimental area and those which collected food from the previously learned feeding station or the nonlearned one. During the testing phase, we observed that a low rate of wasps trained with 2 collecting visits returned to the experimental area (22%), whereas the rate of returning wasps trained with 4 or 8 collecting visits was higher (51% and 41%, respectively). Moreover, wasps trained with 8 feeding visits collected food from the previously learned feeding station at a higher rate than those that did from the nonlearned one. In contrast, wasps trained 2 or 4 times chose both feeding stations at a similar rate. Thus, significantly more wasps returned to the previously learned feeding station after 8 repeated foraging flights but not after only 2 or 4 visits. This is the first report that demonstrates the existence of long‐term spatial memory in V. germanica wasps.     

The influence of past experience on wasp choice related to foraging behavior

Memory has been little studied in social wasps. Vespula germanica （Fab.） （Hymenoptera： Vespidae） frequently revisits nondepleted food sources, making several trips between the resource and the nest. In this study, we analyzed this relocating behavior in order to evaluate whether this species is capable of remembering an established association after 1 h. To this end, we trained wasps to feed from a certain array. Then it was removed, setting it up again 1 h later, but this time 2 baited feeders were put in place, one at the original feeding site and the other opposite the first. We recorded the proportion of returning foragers, and their choice of feeder, after either 1 or 4 feeding trials. After 1 h, 78% of wasps trained with 4 feeding trials and 65% trained with 1, returned to the experimental area. Furthermore, during the testing phase, wasps trained with 4 feeding trials collected food from the previously learned feeder significantly more frequently than from the nonlearned one （P 〈 0.05）. In contrast, wasps that had been trained only once chose both feeders equally. Thus, memory retrieval could be observed 1 h after wasps had collected food on 4 consecutive occasions, but not after only 1. To our knowledge, this is the first study showing that V. germanica is capable of remembering an association 1 h after the last associative event, demonstrating that 1 h does not impair memory retention if4 feeding experiences have occurred.     

Forager success increases with experience inPolybia occidentalis (Hymenoptera: Vespidae)

Summary Foragers of the neotropical swarm-founding waspPolybia occidentalis showed improved task performance, as indicated by foraging success rate, with foraging age. Foragers also spent significantly more time in the field on foraging trips as they aged, while foraging rate did not change with age. These patterns were not explained by directional changes in resource availability or colony need over time. We compare these results to earlier findings on changes in task performance with experience in social insect foragers, and suggest that increases in forager persistence in the field explain improved foraging success with experience.     

Age polyethism and defense in a tropical social wasp (Hymenoptera: Vespidae)

Although tapping the nest in simulated vertebrate attack caused all but 1-day-old workers of Polybia occidentalis to rush from inside and cover the envelope, defense itself—attack of a target—was shown to be carried out only by workers older than about 10 days. While foraging was also performed by older workers, the frequency-age distribution of defensive behavior was skewed to a significantly younger age than was that of foraging. The plot of mean individual probability of defending as a function of age was logistic, leveling off at 0.11 by the age of about 13 days. The distribution of the probability of defending among workers 13 days of age was random, indicating that there is no specialized defender class among older workers. Workers that defended were recruited from among both nest workers and foragers. Although foragers were more likely to defend than were nonforagers, nonforagers contributed more individuals to the pool of defenders because they were more numerous.     

Nest-founding behavior induced in the first descendants ofPolistes dominulus Christ (Hymenoptera: Vespidae) colonies

Summary First descendants' founding ability was tested inP. dominulus colonies by experimentally removing the original nests and queens 21–24 days after the first emergence. Foundation tests were carried out after three different periods had elapsed after the removal and the results were analyzed with regards to the effects of participation in social tasks, duration of stay within the colony and seasonal factors. 1.Immediately after the removal, the foundations that occurred were performed almost entirely by workers, regardless of the duration of their stay within the colony. The number of foundations depended on the season in which the females had emerged. 2.Two months and four months after the removal, foundations were made by only a small number of females (workers and non-workers). These females, which were therefore able to found a nest the year they were born, were mostly among the very first-born individuals in the colonies. 3.The following year after hibernation: females (workers and non-workers) which had stayed for only a short time within the colony (1–15 days) had the highest survival rates. The foundation rates among the surviving females depended only on seasonal factors, since the foundation rates were higher among the non-workers which had emerged later in the season. The results throw some light on the first descendants' nest-founding potential in colonies developing under natural conditions, where many workers in fact stay for only a short time at the nest.     

Effects of experience on food-searching behavior in the antFormica schaufussi (Hymenoptera: Formicidae)

Foragers of the ant Formica schaufussitend to return to and search at a site of a previous food find. The search tactic employed by a forager on its return trip is related to the type of food previously encountered: search is more persistent in response to carbohydrate than to protein food. Using different reinforcement schedules with carbohydrate and protein food rewards, we show that, on a short-term as well as on a long-term basis, the basic pattern of search observed in naive foragers is only slightly modified through foraging experience. Foragers do not increase their search effort or adjust their search pattern when either type of food is systematically renewed on a fixed reward schedule and thus do not seem to be able to learn to assess the food predictability. Collective responses that could compensate for this lack of individual flexibility and increase foraging efficiency at the colony level are discussed.     

Polymorphic microsatellite loci for the Japanese paper wasp,Polistes chinensis antennalis (Hymenoptera: Vespidae)

We isolated 13 microsatellite markers for the Japanese paper wasp Polistes chinensis antennalis. Null alleles were ruled out in the loci because we could detect polymerase chain reaction products for 578 haploid males. Our results indicate that these markers are excellent tools for the analysis of pedigrees within colonies for species with a complicated pedigree structure due to worker reproduction (male production by workers in queen‐right colonies).     

Time allocation of the parasitoidAphidius colemani (Hymenoptera: Aphidiidae) foraging forAphis gossypii (Homoptera: Aphidae) on cucumber leaves

The time allocation of individualAphidius colemani female parasitoids foraging forAphis gossypii nymphs on cucumber leaves has been investigated. Apart from experiences on the current leaf (such as density of hosts on the current leaf, density of hosts on a neighboring leaf and encounters with hosts on the current leaf), the effect of previous leaf visits on the time allocation was studied. Behavioral records were analyzed by means of the proportional hazards model, to determine the tendency of leaving the current leaf. The leaving tendency decreased only on leaves with a high host density (100 aphids), thus increasing the giving up time since the latest encounter. Rejection of aphids had no influence on the leaving tendency. To assess the effect of the number of hosts encountered on the leaving tendency, we considered three classes: 0–30 encounters, 31–100 encounters, and 100 or more encounters with hosts. The effect of the number of hosts encountered differed at different aphid densities. When fewer than 10 aphids were present the leaving tendency was much greater after 30 encounters than beforehand. At a density of 100 aphids the leaving tendency was lower than at the other aphid densities and increased only after 100 encounters. The density of hosts on a neighboring leaf, ranging from 0 to 100 hosts, had a negligible effect on the leaving tendency. Repeated visits to leaves with 10 unparasitized aphids resulted in an increase in the leaving tendency after 10 visits. It is argued that the parasitoids have some innate expectancy of host availability and that they concentrate on high-density patches.