A comparison of nest construction behavior in two species ofPolistes paper wasps (Insecta,Hymenoptera: Vespidae)

Polistes fuscatus variatus (a north temperate wasp) andP. instabilis (a tropical wasp) begin building their nests by applying pulp in a linear sequence of steps: surface preparation, petiole, flat sheet, first cell. After the initiation of first cell there are multiple locations for pulp addition (petiole, new cell, old cell), which become increasingly varied inP. fuscatus as the nest grows (secondary petioles, comb back, substrate, and pupal caps). AlthoughP. fuscatus also adds oral secretion to the petiole, comb back, and pupal caps,P. instabilis only applies oral secretion and no pulp to these locations and does not build secondary petioles or place pulp on the substrate. Both species show a similar series of behavioral acts leading from pulp foraging to application. In both species queens initiate more cells than do subordinates; nonetheless, subordinates account for a large percentage of initiated cells. During pre-emergence and early post-emergence the majority of pulp is used for cell lengthening. BothP. instabilis andP. fuscatus show a bimodal pattern of growth in cell number. The numbers of cells in a mature comb at the end of the nesting cycle are similar in both species (P.f.=274.4 andP.i.=282.6 cells).     

Promotion of thermoregulatory insulation in nests of neotropical wasps by building extra-combs with empty cells

We examined the hypotheses that the empty combs of Polybia occidentalis Olivier (Hymenoptera: Vespidae) nest insulate the inside of the nest. To examine this hypotheses, two kinds of temperature measurements were carried out: 1) with the outer comb of the nest intact (Control) and 2) removed (Treatment), using a large and a small nest. In the large nest, the daily nest temperatures (outer part, Tn1; inner part, Tn2) in Control were lower by 0.6 degrees C (Tn1) and 1.2 degrees C (Tn2) than those in Treatment, because of a higher ambient temperature (Ta) throughout temperature assessment in Treatment. However, the excess temperature (Tn - Ta) in Control was higher than that in Treatment. The value was higher by 0.7 degrees C at Tn1 and 0.1 degrees C at Tn2. In the small nest, the excess temperature in outer part was similar between experiments, while that in inner part of Control was lower than that of Treatment. The temperature fluctuation in the nests was lower in the Control than that in Treatment both the outer and inner part of the comb. We conclude that the unused comb is ecologically invaluable for raising and protecting the brood from extreme changes in ambient temperature during the winter period, because it helps not only keeping a higher nest temperature but also decreasing the temperature fluctuation around the brood combs. In addition, such a high temperature may influence the performance of adult wasps.     

Process of comb division of reused nests in the Australian paper waspRopalidia plebeiana (Hymenoptera,Vespidae)

Summary InRopalidia plebeiana, combs made in the previous year are often reused by foundresses in the following spring and structurally divided into subnests by them. Close observations of the comb dividing process revealed that: 1) combs that had multiple brood areas (areas where cells had eggs) were frequently divided into subnests, while those with single brood areas were never divided; 2) groups of foundresses each occupied a particular brood area even before comb division started; and 3) frequency with which a foundress practiced comb cutting was independent of her social status or oviposition frequency. These observations suggest that the initial grouping of foundresses that colonize an old nest leads to subsequent comb division, rather than the major egg layers cutting combs to safeguard their oviposition territories from their rivals.     

Observations onApoica pallens,a nocturnal neotropical social wasp (Hymenoptera: Vespidae,Polistinae, Epiponini)

Summary Colonies and nests ofApoica pallens in the llanos region of Venezuela range from small foundress nests to large mature colonies. Nests are sited on small diameter, near-horizontal branches in a variety of shrub and tree species. During the day, adult wasps cluster on the face of the nest in an array that seems to be determined by orientation to gravity; defense of the colony against parasitoids and ants by the resting wasps may be more a passive than an active behavior. Wasps fan their wings to cool the colony during the day, but no foraging for water accompanies the fanning behavior. Nightly foraging activity begins with the explosive departure from the nest of hundreds of wasps, most of which rapidly return. Moderate foraging levels early at night give way to very low foraging levels in pre-dawn hours. The period of moderate foraging may be extended for longer hours during increased moonlight. Foraging wasps collect arthropod provisions for larvae. Larvae produce a trophallactic saliva; adults engage in inter-adult trophallaxis; brood are cannibalized. During cluster formation prior to swarm emigration, adult wasps do not appear to scent-mark substrates such as leaves. Instead,A. pallens exhibits a calling behavior, unique among polistine wasps studied to date, in which the gaster is held rigidly away from the thorax and metasomal sternal glands are exposed. Swarms can emigrate during the day.A. pallens may incorporate absconding and colony relocation as features of its colony cycle in the highly seasonal llanos.     

Home Range Analysis in the Study of Spatial Organization on the Comb in the Paper Wasp Polistes Dominulus

In order to understand how the apparent freedom of individual movements can concur with social order in a colony, we investigated spatial relationships between foundresses, workers and immature brood in a paper wasp. This is the first time an ecological analytical approach (home range analysis) has been employed to describe small scale spatial use in a social context. In this study, home range was not used in its strictly ecological sense, but rather applied to the comb. Our results show that the positions individual wasps assumed on Polistes dominulus combs are spatially structured. Workers and foundresses do not occupy the comb in random distribution, but rather follow predictable spatial patterns. Each active wasp has its own spatial fidelity area, at least over a 1 d observation period, spending the majority of its time within a very small area averaging approx. 12% of the comb but occasionally it may cover an area of up to 50% of the comb surface. Dominant females occupied a significantly smaller area that either subordinate foundresses or workers. Areas patrolled by workers varied in shape and size, with no relation to time spent on the comb, wasp density or position of immature brood. All the wasps clumped around the dominant female, who proved to be the only individual affected by cell content. Nevertheless, although the wasps tended to clump together, average superimposition rates among nest‐mates was very low (higher in workers than foundresses), suggesting that the wasps limit each other’s individual spaces.     

Ground-nesting bees determine the location of their nest relative to a landmark by other than angular size cues

Bees and wasps acquire a visual representation of their nest's environment and use it to locate their nest when they return from foraging trips. This representation contains among other features cues to the distance of near-by landmarks. We worked with two species of ground-nesting bees, Lasioglossum malachurum (Hymenoptera: Halictidae), Dasypoda hirtipes (Hymenoptera: Melittidae) and asked which cues to landmark distance they use during homing. Bees learned to associate a single cylindrical landmark with their nest's location. We subsequently tested returning bees with landmarks of different sizes and thus introduced large discrepancies between the angular size of the landmark as seen from the nest during training and its distance from the nest. The bees' search behaviour and their choice of dummy nest entrances show that both species of ground-nesting bees consistently search for their nest at the learned distance from landmarks. The influence of the apparent size of landmarks on the bees' search and choice behaviour is comparatively weak. We suggest that the bees exploit cues derived from the apparent speed of the landmark's image at their retina for distance evaluation.     

Organization and Regulation of Nest Construction Behavior in Metapolybia Wasps

Field observations and experiments revealed that construction behavior of Metapolybia wasps is based on parallel processing and distributed decision making. Sixteen behaviors were used to separate five behavioral groupings: specialized water forager, flexible pulpforager, active builder, active generalist, and idle. The idle category proved to be the source and the sink of the other task groups, although specialist foragers tend to retain their duties or take over other active roles. Nest construction is partitioned into three tasks. Pulp foragers transfer wood-pulp to the nest where other wasps (builders) distribute and process it further. The builders incorporate this material into the nest structure on the basis of individual decisions. Water foragers provide the extra water necessary for both building and pulp collecting. Material exchange takes place on the nest between pairs or in small groups. The duration and frequency of different behaviors, the number of wasps belonging to different behavioral groups, and the different scale of specialization in different groups suggest that the colony-level performance and speed are governed by the activity of the pulp foragers, who receive information about both the water saturation level of the colony and the activity of the builders through local interactions. Several predictions of this hypothesis were supported by disturbing the normal construction behavior through removing or decreasing the number of individuals belonging to different behavioral groups or supplying additional building material.     

Scaling in nests of a social wasp: a property of the social group

The numbers of brood cells in nests built by founding swarms of the Neotropical social wasp Polybia occidentalis closely correlate with the numbers of wasps in the swarms. We analyzed nests of different sizes to determine how they scale with respect to the allocation of brood cells among combs. Three patterns were evident: compared to smaller nests, larger nests have (1) more combs and (2) larger combs; and (3) among nests containing the same number of combs, the last two combs diverge in relative size as nest size increases. Taken together, these results suggest that members of a swarm somehow "know" the size of the swarm they are in. This information feeds back to individual builders, which quantitatively modulate their responses to stigmergic cues in ways that result in the nest-size-scaled allocation of brood cells among combs. The patterns also suggest that swarms fine-tune the final size of their nests by making corrections as they build.     

Factors affecting the acceptance of alien conspecifics on nests of the primitively eusocial wasp,Ropalidia marginata (Hymenoptera: Vespidae)

In the primitively eusocial wasp, Ropalidia marginata,individual females are known to drift from one newly founded nest to another. In the laboratory, young (<6- to 8-day-old) alien wasps are accepted onto unrelated colonies, while older (>6- to 8-day-old) wasps are not. Here we have investigated the factors that could influence the acceptance of foreign conspecifics onto unrelated nests. Individually marked wasps of different ages, isolated immediately after eclosion from the natal nest and from each other, were introduced onto unrelated recipient nests. Considered separately, both age and ovarian condition seemed to influence the probability of acceptance as well as the levels of aggression and tolerance received by the introduced wasps. However, partial correlation analysis and multiple regression analysis indicated that only age had a direct influence and that the ovarian condition acts only through age, a variable with which it is highly correlated. The observed acceptance of young aliens and rejection of old aliens are less likely to be due to the perception of older wasps as a reproductive threat rather than some age-related factor, other than ovarian condition, for example, the relative ease with which younger wasps can be molded into desired roles.     

Pulp partitioning and worker specialization in polistine wasps (Hymenoptera,Vespidae, Polistinae)

The character of pulp partitioning was studied in three Palaearctic polistine wasps (Polistes dominula and P. gallicus in the Black Sea Biosphere Reserve, and Polistes nimpha in the Lugansk Nature Reserve, Ukraine) and in five Neotropical species (P. lanio, P. versicolor, Mischocyttarus injucundus, M. alfkeni, Metapolybia cingulata, and Protopolybia exigua in Trinidad). In independent-founding Polistes and Mischocyttarus species, all cases of pulp partitioning were connected with aggression on the part of the foundresses. An experimental increase of nesting density of P. dominula led to changes in the relative abundance of workers in the polyethic task groups. Some workers practically stopped hunting and switched over to pulp delivery; some of these workers possessed well-developed ovaries and were able to lay eggs. The experimentally removed part of the nest envelope in the swarm-founding wasp Metapolybia cingulata was restored only in colonies that included young workers. The specific features of nest building activity in independent-founding and swarm-founding polistine wasps are discussed.     

Interaction between the solitary bee Chelostoma florisomne and its nest parasite Sapyga clavicornis − empty cells reduce the impact of parasites

1. Nesting behaviour and interactions between the bee Chelostoma florisomne (L.) (Megachilidae) and its nest parasite Sapyga clavicornis (L.) (Sapygidae) were studied through continual observations of individuals and dissections of bee nests. Protection of bee offspring is based on (1) the bee’s discovery and removal of parasite eggs deposited prior to the construction of a cell closure, (2) minimising the time when fully provisioned cells might be parasitised successfully, and (3) the construction of empty cells in front of brood cells. 2. An empty cell was found in front of 64.4% of all brood cells and, if the outermost brood cell in a nest was excluded, in front of 74.3% of inner brood cells. A vestibule closure is most often constructed in front of the outermost brood cell. 3. Following oviposition, the bee made only five flights, which together lasted 6–13 min, to construct a cell closure. A cell closure does not prevent the nest parasite from oviposition inside the brood cell, however, and parasite eggs deposited through the cell closure are not detected and removed by the bee. Only an additional cell closure, i.e. the formation of an empty cell, may protect a brood cell when the bee is not in the nest. The nest parasite often oviposited through the additional cell closure but its offspring were then trapped in the empty cell and starved to death. 4. Only 5.4% of the inner brood cells that were protected by an empty cell were parasitised, compared with 28.9% of those without an anterior, empty cell; 27.4% of the empty cells contained dead parasite offspring (eggs and larvae). Thus, the empty cells provided significant protection and, combined with additional means of protection of brood cells, led to a low degree of parasitism. More than 77% of the wasp offspring died at an early stage due to intraspecific interference competition within brood cells and as result of the wasps’ oviposition into empty cells.     

Nesting biology of four species of Trypoxylon (Trypargilum) (Hymenoptera: Crabronidae) in Chaco Serrano woodland, Central Argentina

Trypoxylon (Trypargilum) (Crabronidae) wasps are solitary spider predators that can build their nests in artificial trap-nests, which enables study of their nesting architecture and biology. Twenty traps (each containing 15-30 internodes of cane) were placed in each of nine sites of Chaco Serrano Woodland in Central Argentina (Córdoba) in October 2005, and were recovered in June 2006. We obtained 91 nests of four species of Trypoxylon (Trypargilum). In the laboratory, each Trypoxylon nest was sectioned longitudinally to study its architecture. The number of brood cells was counted, and the occurrence and length of vestibular and intercalary cells and the pre-closing plug space were recorded. We measured the diameter of the cane entrance, total length of the nest, length of each brood cell, maximum thickness of mud partitions and closing plug thickness. We also recorded the cell contents: the wasps, their natural enemies and the prey spiders. Mortality was assessed and the sex ratio calculated for each species. Finally, the nests were examined to help clarify the function of the vestibular cell. The nest architecture was similar in the four species, with linear brood cells located one after the other separated by mud partitions, as in other species of the subgenus Trypargilum. Forty-eight percent of the nests had vestibular cells, but only two had intercalary cells. The thickness of the mud partitions and the length of the brood cells differed among species and were related to the size of the emerged wasp. The diameter of the nest entrance was directly related to the average length of the fore-tibia. Sex ratios of all species did not deviate from 0.5. Mortality due to parasitoids (Eulophidae; Melittobia sp.) was similar among species, while the mortality due to cuckoo wasps (Chrysididae) in T. lactitarse was higher than in the other species. The presence of vestibular cells was not related either to the mortality due to natural enemies or to the orientation of the trap in the field. Spiders in the family Araneidae were the most frequently collected prey.     

Allocation of Colony‐Level Foraging Effort in Vespula germanica in Response to Food Resource Quantity,Quality, and Associated Olfactory Cues

In social insects, selection takes place primarily at the level of the colony. Therefore, unlike solitary insects, social species are expected to forage at rates that maximize colony fitness rather than individual fitness. Workers can increase the net benefit of foraging by responding to increased resource availability, by responding more strongly to higher‐quality resources, and by decreasing the uncertainty with which nestmates find resources. Unlike many ants and social bees, no social wasp is known to utilize a nest‐based recruitment signal to inform nestmates of food location. On the other hand, wasps do learn the odor of food brought to the nest and use this cue to locate the food source outside the nest. Here, we quantify the effects of three food‐associated variables on the allocation of foraging effort in the yellowjacket Vespula germanica. We used an experimental approach to assess whether resource quantity, quality, or associated olfactory information affect the probability that a forager will leave the nest on a foraging trip. We addressed these questions by inserting a known amount of sucrose solution directly into nests and recording foraging effort (departure rate) over the subsequent hour‐long observation period. No differences were found in foraging effort because of the presence/absence of olfactory cues, but there was strong evidence that foraging effort increased in response to resource influx and resource quality. Thus, while olfactory cues are learned in the nest, only resource quality and the cue of increased amount of food in the nest factor into a forager's decision of whether or not to depart on a foraging trip. However, as prior work has shown, once a wasp forager leaves the nest, it uses the learned olfactory cues to aid in finding resources.     

Nest surface hydrocarbons facilitate nestmate recognition for the social wasp,Polistes metricus Say (Hymenoptera: Vespidae)

Nestmate recognition was observed inPolistes metricus Say workers only if the workers were exposed to their nest surface hydrocarbons after eclosion. If the newly emerged workers were never exposed to the nest hydrocarbons as adults, they showed no discrimination between nestmates and nonnestmates. Furthermore, the newly emerged workers were accepted more readily by their experienced nestmates than by experienced nonnestmates only if the newly emerged wasps had been exposed to the nest surface hydrocarbons. This reciprocal recognition implies that the nest recognition cues are nest surface hydrocarbons that are learned and that may be acquired byP. metricus workers as adults on the natal nest.     

Potential consequences of angling on nest-site fidelity in largemouth bass

Breeding site fidelity has evolved in many vertebrate taxa, suggesting both that site selection has an important influence on fitness potential and that the decision to reuse a nesting site is related to the individual’s prior nesting success at that location. For a species that provides parental care, such as the Largemouth Bass Micropterus salmoides, catch-and-release angling impacts individual nesting success and fitness through physiological disturbance and by removing the nest-guarding male from its brood, thereby allowing temporary access to eggs and hatchlings by brood predators. To assess the impact of catch-and-release angling on nest site fidelity, we compared the consequences of angling on individually marked (i.e., with passive integrated transponders) nest-guarding male Largemouth Bass in Ontario. An extremely high degree of nest site fidelity in year two was observed for males that were angled only once during year one (87% within 10 m of the previous year’s nest), 96.7% of which remained on the nest and completed parental care activities. There was significantly lower fidelity in year two, however, for males that were angled multiple times during year one (27% within 10m of the previous year’s nest), only 5.6% of which remained on the nest and completed parental care activities. This observed difference suggests that angling nesting bass may cause them to avoid previously used nest sites and instead search for alternative sites during future reproductive seasons. This human-induced impact on nest site choice may impact the future reproductive success of those Largemouth Bass.     

Colony Membership, Division of Labor, and Genetic Relatedness Among Females of Colonies of Eustenogaster fraterna (Hymenoptera, Vespidae, Stenogastrinae)

To compare the social structure of primitively social wasps with that of communal breeding vertebrates, we used a new technique based on micro videocameras applied to the nest envelope to study the organization of behavior in Malaysian colonies of the stenogastrine wasp Eustenogaster fraterna. The reproductive division of labor in this species appears to be different from that reported so far in other species of Stenogastrinae: it is at a very primitive stage because the helpers work less hard and take fewer risks than the egg-layers. Nevertheless, the very low-risk tasks performed by the helpers (nest guarding and supply of abdominal secretion to the young larvae) are important for the colonies. Behavioral characters and ovarian development of the helpers suggest that females are only temporary helpers and that they wait to start their own production of offspring in the hope of inheriting the nest. Females of the same colony can be highly related, but in some colonies we found low-related individuals. The social organization of these wasps resembles that of vertebrate groups with helpers at the den, i.e., in the small number of individuals, the division of labor with limited reproductive inhibition, and the prolonged external work of the reproductive individuals. However, the main drive to sociality in these insects appears to be different, being based mainly on problems connected with short adult life and long immature brood development.     

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.     

Behavioral role differentiation in the primitively eusocial waspBelonogaster juncea juncea (Hymenoptera: Vespidae)

With a view toward describing behavioral variability among individuals of the primitively eusocial speciesBelonogaster juncea juncea, we recorded the time-activity budget spent on five behavioral categories (foraging, building, feeding, inactivity, and reproduction) by 52 individuals belonging to four postemergence colonies. A principal-components analysis coupled with a hierarchical cluster analysis enabled us to discern four behavioral roles. The reproductive role is reserved to one individual (functional monogyny) and the workers’ role is differentiated into foragers, builders, and guards. The foragers are females that spend the most time (82.6% of their time) foraging, supplying the nest with prey load and liquid matter. The builders are individuals (with 41.5% of their time off the nest) that tend to bring pulp into the nest and then undertake building activities. The guards are those females that spend the most time (79.7% of their time) being inactive on the nest.     

Cooperative nest building and brood care by nestmates and non nestmates in Ropalidia marginata : implications for the evolution of eusociality

In the primitively eusocial wasp, Ropalidia marginata, low levels of intra-colony genetic relatedness, lack of intra-colony kin discrimination and acceptance of young wasps into alien colonies, prompted us to investigate whether or not there exists a cost of such high genetic variability. Freshly eclosed wasps were paired either with their nestmates or with their non nestmates and their performance in nest building and brood care were compared. There was no demonstrable difference between nestmate and non nestmate pairs in terms of success in raising adult offspring, time required for nest initiation, brood developmental period and productivity. There was also no difference in the efficiency of cooperation and division of labour between the nestmate pairs and non nestmate pairs. These results reinforce the idea that the haplodiploidy hypothesis is insufficient to explain the prevalence of worker behaviour in R. marginata and emphasize the importance of factors other than genetic relatedness in the evolution of eusociality. Received: 27 April 1998 / Accepted: 10 July 1998     

Nesting and biology of Jucancistrocerus caspicus (hymenoptera,vespidae, eumeninae)

Females of Jucancistrocerus caspicus nest in dense clay ground on the vertical surface of cliffs. The nests contain 1–9 cells (on average 2.8) and have a linear-branched construction. Females surmount the entrance of the burrow with a curved chimney which has a laced structure. The cells are positioned vertically in the main burrow and obliquely or horizontally in the lateral tunnels; the cells in a row are separated with double partitions. The size of the cells is 7–9 × 4–4.5 mm, the diameter of the nest burrow is 4 mm. The egg is laid before provisioning and is attached to the cell ceiling with a filament. Females hunt for weevil larvae and store 23–33 larvae (on average 27.8) in each cell. The species is univoltine, with prepupae hibernating in their cocoons. The nests are parasitized by the cuckoo wasps Chrysis rutilans which cause 11.5% of brood mortality. Adult wasps are killed by the spiders Pholcus sp. living near the nests.