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.     

Distinguishing signals and cues: bumblebees use general footprints to generate adaptive behaviour at flowers and nest

Chemicals used in communication are divided into signals and cues. Signals are moulded by natural selection to carry specific meanings in specific contexts. Cues, on the other hand, have not been moulded by natural selection to carry specific information for intended receivers. Distinguishing between these two modes of information transfer is difficult when animals do not perform obvious secretion behaviours. Although a number of insects have been suspected of leaving cues at food sites and nest entrances, studies have not attempted to experimentally distinguish between cues and signals. Here, we examine the chemical composition of the scent marks left by the bumblebee Bombus terrestris at food sites and compare it to those found at a neutral location. If bees are depositing a cue, we expect the same chemicals to be found at both sites, but if they deposit a signal we only expect to find the scent marks at the food site. We were also interested in identifying the chemicals left at the nest entrance to determine if they differed from those used to mark food sites. We find that bees deposit the same chemicals at food, nest and neutral sites. Therefore, bumblebees leave behind general chemical footprints everywhere they walk and we propose that they learn to use these footprints in a manner that ultimately enhances their fitness, for example, to improve their foraging efficiency and locate their nest. Experimentally, distinguishing these two modes of information transfer is crucial for understanding how they interact to shape animal behaviour and what chemical bouquets are under natural selection. Handling Editor: Heikki Hokkanen     

Landmark orientation during the approach to the nest in the stingless beeTrigona (Tetragonisca) angustula (Apidae,Meliponinae)

Summary We displaced a small nest box containing stingless bees (Trigona (Tetragonisca)angustula) over distances of up to 1.6 meters in different directions and counted the numbers of returning foragers to measure the effects of this manipulation on the homing ability of bees. Bees find it hard to locate the nest box when it was displaced more than about 1 m backwards, forwards or sideways relative to the direction into which the nest entrance pointed. They do not find the nest when its height above ground is changed. The bees use landmarks in the vicinity of the nest to locate it: When the nest box is displaced and landmark positions are changed so that their angular position at the new nest site is the same as at the normal nest position their homing ability is less impaired than it is without changes in landmark positions. Our results show that the bees do not use the nest box itself as a landmark until they have approached the nest position to within about 1 meter with the aid of surrounding landmarks.     

Not all who wander are lost: nest fidelity in <Emphasis Type="Italic">Xylocopa virginica</Emphasis> examined by mark recapture

Relocation to novel nests (sometimes called drifting) in flying Hymenoptera is often interpreted as the result of navigation error and guard bees erroneously admitting foreign individuals into the nest. We studied nest fidelity and nest relocation of both females and males in a nesting aggregation of Xylocopa virginica in southern Ontario, Canada, where females can nest either solitarily or socially. Adult female and male bees were trapped at nest entrances, individually paint marked, and then released. Subsequent recapture patterns were used to assess nest fidelity: that is, how faithful individuals were to their home nest and how often they moved to another nest. Bees were considered to have relocated if they were recaptured in a nest different from the one in which they were initially trapped, indicating that they had spent at least one night in a new nest. Some females were only captured in one nest, some occasionally moved to new nests, temporarily or permanently, and a few were never caught in the same nest twice. In addition, females relocated to nests that were further away in 2007 when population density was low, suggesting that they seek out and claim nesting spaces when they are available. Males relocated more frequently than females, with most drifting from nest to nest in no obvious pattern. This indicates that males spend the night wherever space is available or in nests nearest to their territories. This study reveals that for both female and male X. virginica, nest membership is not as stable as once thought.     

Visual marks learned by the solitary bee Megachile rotundata for localizing its nest

We studied homing behaviour of leaf-cutter bees, Megachile rotundata, by using artificial landmarks. We evaluated their nest-searching behaviour in different test situations to elucidate the nature of the visual marks they used in this task. When we modified or removed geometrical figures surrounding the nest, the bees searched for longer, showing that they noticed the introduced changes. However, these manipulations never prevented bees from finding their nest, suggesting that other visual cues were crucial in the task. Manipulations of the edges provided by the boundaries of the device (nest block, metal sheet on which the block was mounted) strongly impaired the homing performance. The further away the edges that were left intact, the stronger was the impairment of the homing behaviour. These results suggest that bees learn the distances of the various edges from the goal and that edges have a hierarchical significance according to their distance from the nest. The most distant edges provide vague information, which suffices to guide the insect towards the next edge in the sequence, until it recognizes the final, precise location of the nest. The results support the conclusion that information on distances is acquired using cues derived from motion parallax generated by the insect's self-motion. Recognition of edge parameters such as position and orientation might be achieved by an image-matching mechanism based on dynamic processes. Thus, in the homing task, there is no clear discrepancy between the eidetic and the parametric hypotheses of spatial representation.     

Floating and fidelity in nest visitation byCrawfordapis luctuosa (Hymenoptera: Colletidae)

We studied nest visitation behavior of the ground-nesting beeCrawfordapis luctuosa (Colletidae) at the Monteverde Cloud Forest Preserve, Puntarenas Province, Costa Rica. Some females entered many nests, while others consistently entered only one nest. Multiple nest entry was exhibited by bees we called floaters, and those bees that consistently entered only one nest were provisioning bees. Provisioning bees abandoned their old nests after provisioning periods and became nonprovisioning bees that floated over the nesting area. Floating behavior was a nest-searching behavior typified by oscillating flights just above the ground (<15 cm). Floaters investigated available nests and were not observed to interact aggressively with nest owners. Upon establishing a new nest, floaters became provisioning bees.     

How Do Carpenter Bees Recognize the Entrance of their Nests?: An Experimental Investigation in a Natural Habitat1

Carpenter bee females (genus Xylocopa) enter only their own nests. They can be found in the same burrow over a period of months, regardless of whether they arc living singly or aggregated in close vicinity. The aim of this study was to investigate and to clarify which sensory systems are responsible for the recognition of the nest. Females of three different African species were tested in their natural habitat. Following alteration in the immediate visual surroundings of either the entrance or the entire dwelling stem, bees behaved as they did in undisturbed situations and entered their nests. Furthermore, a relative dislocation of the entrance within the dwelling stem or a displacement of the whole stem within a restricted range had no effect on the recognition of the burrow. When the entrance was plugged by foam rubber the bees landed only after extended searching. When the entrance was closed by a strip of scotch tape the bees searched for several minutes, departed, then returned and searched again with the same result. The bees showed no indications that they recognized their burrow. These results strongly support the conclusion that the bees recognized their burrows primarily or exclusively by olfactory cues, and, furthermore, that individual odors were involved. The possible contributions of different sensory systems for the recognition of the nest and the possible biological function of the use of olfactory cues are discussed.     

Nest Entrance Marking with Colony Specific Odors by the Bumble Bee Bombus occidentalis (Hymenoptera: Apidae)

We investigated the ability of naturally foraging female bumble bees (Bombus occidentalis) to recognize their nest entrance in the field. In a blind binary-choice paradigm, females discriminated between nest entrances tipped with caps from their own nest entrances and entrances tipped with caps previously associated with foreign heterospecific or foreign conspecific colonies. Washing caps in organic solvents eliminated the manifestation of this nest-entrance recognition and significantly decreased the episodes of antennating caps. These results indicate that bees deposit colony-specific chemicals on nest entrances that are later perceived via contact chemoreception. Females treated foreign heterospecific and foreign conspecific caps similarly, suggesting that bees may not assess or respond to the degree of similarity between their own colony odor and a foreign colony odor.     

The Significance of Odor Cues and Dance Language Information for the Food Search Behavior of Honeybees (Hymenoptera: Apidae)

Although several independent lines of evidence show that bees can make use of information provided by their dance language, there is an ongoing controversy about the significance of the dance information versus odor cues in the search behavior of recruited bees. A series of experiments was performed to assess the relative significance of dance information and odors for the site-specific search of recruit bees. In these experiments recruit bees were trapped automatically at arrays of artificial flowers at various distances from the hive. The distribution of directions in which the recruits searched for food was compared between recruitment by dancers performing well-oriented dances on the vertical side of the comb and dancers performing disoriented dances on a horizontal comb. The results show quantitatively that bees use both odor cues and the dance information. The greater the distance to the feeding site, the greater is the relative significance of the dance information.     

The relocation of ant nest entrances: Potential consequences for ant-dispersed seeds

The rate at which ant nests relocate may affect the fate of ant-dispersed seeds by influencing nutrient and seed accumulation in localized areas. In this study, the movement of ant nest entrances was monitored in fixed quadrats in dry sclerophyll woodland in Kuringai Chase National Park, NSW. Changes in entrance location were rapid for most species, with few entrances remaining in use for more than 6 months. Approximately 30% of entrances that closed were subsequently re-opened. There was no obvious seasonal pattern in entrance relocation. After 1 year, between 5 and 40% of the ground surface of the quadrats had been within 10 cm of a nest entrance. New nest entrances did not cluster near old entrances, indicating that nest entrance relocation may be accompanied by changes in underlying nest structure. Nutrient levels in soil samples from active nests of Rhytidoponera metallica, Aphaenogaster longiceps, Pheidole sp. 1 and Iridomyrmex sp. 8 did not differ significantly from random locations. The rapid changes in entrance location and the lack of nutrient enrichment may be the result of continual and progressive underground shifts in nest location. Such shifts have three potential consequences for seeds that remain buried within the nests. First, seeds will not benefit following germination by being in a nutrient-rich microsite. Second, a proportion of seeds collected by harvester ant species may escape predation if left in an abandoned section of the nest. Third, concentrations of seeds in localized areas may be reduced, leading to a reduction in competition between establishing seedlings.     

Intra-and intercolony patterns of nest dispersion in the ant Lasius neoniger: correlations with territoriality and foraging ecology

Colonies of the ant Lasius neoniger have multiple nest entrances that are distributed throughout a colony's foraging area. Associated with each nest entrance is a group of workers that show strong fidelity to that nest entrance. Territorial expansion, as indicated by increases in the number of nest entrances per colony, is correlated with foraging activity. Although there is variation between colonies in the seasonal pattern of territorial expansion, most nests become active in early summer, increase the size of the area foraged until midsummer, and then decrease the number of active nest entrances in late summer. Over the study plot as a whole, the dispersion pattern of nest entrances changed from clumped, or tending to be clumped, in early spring to random in mid-and late summer. Within colonies, nest entrances were significantly overdispersed. Intra-and interspecific competition negatively affected foraging, and workers from a given nest entrance were most successful at retrieving prey less than approximately 15–20 cm from the entrance. The average distance between nest entrances within a colony was 37.7±3.3 cm (mean±95% confidence interval, n=115), which is approximately twice the distance at which workers can retrieve prey. The polydomous nest structure of L. neoniger appears to partition territory within a colony by spatial subdivision of its foragers, and thus may reduce loss of prey to competitors.     

Rules for Nest Sanitation in a Social Spider Mite, Schizotetranychus miscanthi Saito (Acari: Tetranychidae)

Waste management behavior is essential to achieve nest sanitation that is highly inferential on the evolution of group living because nest waste is an inevitable cost. However, how group living animals dispose of waste has not attracted much attention. Schizotetranychus miscanthi Saito is a social spider mite infesting a perennial grass (Miscanthus sinensis Anderss), in which all nest members tend to defecate at specific sites. We investigated the mechanisms by which the individuals select the site of defecation. The results show that nest members defecate at only one site inside the nest, and that waste management is maintained by two simple rules. First, mites defecate near the nest entrances if no volatile chemical cues are available, and secondly, when chemical cues are available from feces deposited previously, they defecate at this site. We discuss the adaptive significance of these mechanisms, as well as their role in the evolution of sociality in mites.     

Wilson's Storm Petrels Oceanites oceanicus Recognise the Olfactory Signature of Their Mate

Chemical signals in birds have rarely been considered as recognition cues. Nevertheless, recent experiments showed that several petrel species are able to recognize their nest by smell, and in at least one species even their mate. But the use of smell may be different across the petrel species and olfactory nest recognition appears to be dependent on species’ breeding biology. To increase our knowledge of individual olfactory recognition in petrels and the relationships between breeding biology and use of smell, we tested Wilson’s storm petrels Oceanites oceanicus in Antarctica. In previous experiments, these birds failed to home if rendered anosmic, but the method employed to obtain anosmia (potentially stressing birds) and the fact that they breed in 24‐h daylight suggest that they might use visual, rather than olfactory, cues to recognize their nest. Our birds were tested in T‐maze experiments where nest odours or partner odours were presented. Wilson’s storm petrels preferred odours of their own nest and mate. Results on olfactory nest recognition confirm and complete previous results, viz. anosmic Wilson’s storm petrels do not home. Storm petrels olfactory mate recognition suggests that this ability may be widespread in burrowing petrels and implements olfactory nest recognition.     

Biology of <Emphasis Type="Italic">Lasioglossum</Emphasis> (<Emphasis Type="Italic">L.</Emphasis>) <Emphasis Type="Italic">majus</Emphasis> (Hymenoptera: Halictidae), a largely solitary sweat bee with behavioural adaptations to communality

Within the bee family Halictidae there have been three origins of sociality. Although detection of origins and reversal from sociality requires phylogenetic studies, at a behavioural level a predisposition to sociality can be detected by analysis of intra-specific interactions. We studied aspects of nesting biology and behavioural interactions in Lasioglossum (Lasioglossum) majus, a poorly known halictine inhabiting temperate regions of Europe, which is suspected to be solitary. Nests were found to be largely used by one female, but some were shared by more than one individual. These few nests, whose entrances were very close to each other, were found to be connected underground. A few individuals were observed to enter in a nest where a female was waiting, behaving as a guard and allowing the incoming bee to enter the nest. By use of circle-tube experiments, the behavioural repertoire exhibited by females during encounters was assessed. Levels of withdrawal and cooperative events were comparable with those observed in other solitary nesting species, but aggressive events were very rare, as in several observed communal species. We conclude that L. (L.) majus females, despite general solitary nesting, possess behavioural components enabling them to adopt, probably in high nest-density areas, nest-sharing strategies. A similar kind of local social polymorphism has been observed in two other species of the subgenus Lasioglossum, but these are the first data available on a European species and the first record of subterranean connections among nests of halictid bees.     

PVC nest boxes are less at risk of occupancy by feral honey bees than timber nest boxes and natural hollows

Feral European Honey Bee (Apis mellifera) has been identified as a potential nest competitor for Australian hollow nesting species, but few studies have investigated the impact of feral honey bee competition on Threatened species. Our study used data from Glossy Black‐cockatoo (Calyptorhynchus lathami halmaturinus) nests on Kangaroo Island, monitored and managed over an 11‐year period, and found 12% of nests became occupied by feral honey bees during that period. Our results indicate that feral honey bees were less likely to occupy nest boxes made of PVC (5%) compared with wooden nest boxes (24%) or natural hollows in Eucalyptus trees (14%). The removal of feral honey bee hives from nests is a priority for long‐term conservation of glossy black‐cockatoos on Kangaroo Island. We recommend that PVC nest boxes are chosen for future nesting habitat restoration, due to the more frequent use of wooden nest boxes by feral honey bees.     

Old comb for nesting site recognition by <Emphasis Type="Italic">Apis dorsata?</Emphasis> Field experiments in China

The Asian giant honey bee, Apis dorsata, often conducts seasonal, long-distance migrations in southern China, between a preferred tree (having more than one nest) and alternate sites. Although worker bees cannot make a round-trip journey, colonies re-utilize preferred trees after an absence of several months. We performed comb experiments in which bases and all abandoned combs were entirely scraped off trees and their sites covered with plastic, or comb was moved to trees of the same species. Swarms of giant honey bees investigated trees where combs were removed and continued to nest on the same trees. In contrast, placing combs in nets on previously used trees, or on nearby trees of the same species, did not attract more swarms. The same number of colonies that left them returned to previously occupied trees. Our findings suggest that direct olfactory or sensory contact with old comb bases might regulate nest establishment, but individual trees, lacking normal visual or chemical cues of old nests, are relocated using behavioral devices that remain to be elucidated. Received 12 February 2007; revised 5 June 2007; accepted 13 September 2007.     

Evidence of Entrance Sharing and Subterranean Connections in Andrena agilissima (Hymenoptera; Andrenidae)

Andrena agilissima is a bee that nests on sloped or vertical earth walls. Data from 3 years of fieldwork presented here show that this species is communal and uses the same nesting site for many years. On an earth wall on Isola d'Elba (Tuscany, Italy), covered with many bee nest entrances, we studied an aggregation of Andrena agilissima to determine if bees share entrances, if the sharing provides evidence for subterranean connections, and if bees involved in sharing show a division of labor. We marked several females coming out of the same entrance, as well as recapturing the same females using different entrances. No division of labor was observed in our observations.     

Use of terrestrial chemical trails for nest orientation in an open nesting wasp, <Emphasis Type="Italic">Dolichovespula saxonica</Emphasis> F.

Summary The long-cheeked wasp Dolichovespula saxonica typically constructs exposed nests which can be reached by flying. Usually foragers do not walk on substrates in the close vicinity of the nests as cavity breeding wasps do (Steinmetz et al., 2002). Unexpectedly, when forced to walk outside the nest instead of flying in an artificial tunnel system, D. saxonica foragers lay a terrestrial trail and use it for orientation in the nest area in our experiments. 41% of the foragers followed the trail in a direction they were not accustomed to. We suggest that the foragers have employed the same orientation cues normally used for orientation in the close vicinity of the nest when approaching a free-hanging nest by flying, for example nest odour. Nest odour substances may have been transferred to the substrate as a trail as a consequence of foragers walking through the tunnels.Received 23 August 2002; revised 27 January 2003; accepted 30 April 2003.     

Information analysis by the paper wasp,Polistes fuscatus,during nest construction (Hymenoptera,Vespidae)

Summary The building decision process of the paper wasp,Polistes fuscatus, was studied by 1) analyzing the search pattern of the wasps prior to the addition of pulp to different areas of the nest, 2) comparing the pulp addition needs of the cell chosen for lengthening to those of other cells in the nest, and 3) presenting the wasps with eight types of dichotomous building choices, which provided information about the relative influence of different building cues. Wasps conduct a hierarchical search prior to pulp addition, which means that they search the comb face and petiole disproportionately more often and more thoroughly than the comb back and sides. Once a particular nest area triggers closer scrutiny, comparisons are made with adjoining areas. The most needy location is then chosen based on nest cues. When lengthening a cell, the development of the brood and relative cell length have a strong influence on which cell is chosen at all times, while distance of the brood from the cell mouth becomes important during the later stages of brood development. The results indicate that there is no simple hierarchical weighting of cues. The decision process involves comparisons among multiple cues, which for the most part have an additive influence when variation in relative cue strength is considered.     

A stingless bee (Melipona seminigra) uses optic flow to estimate flight distances

Foragers of a stingless bee, Melipona seminigra, are able to use the optic flow experienced en route to estimate flight distance. After training the bees to collect food inside a flight tunnel with black-and-white stripes covering the side walls and the floor, their search behavior was observed in tunnels lacking a reward. Like honeybees, the bees accurately estimated the distance to the previously offered food source as seen from the sections of the tunnel where they turned around in search of the food. Changing the visual flow by decreasing the width of the flight tunnel resulted in the underestimation of the distance flown. The removal of image motion cues either in the ventral or lateral field of view reduced the bees' ability to gauge distances. When the feeder inside the tunnel was displaced together with the bees feeding on it while preventing the bee from seeing any image motion during the displacement the bees experienced different distances on their way to the food source and during their return to the nest. In the subsequent test the bees searched for the food predominantly at the distance associated with their return flight.