Eastern bluebirds Sialia sialis do not avoid nest boxes with chemical cues from two common nest predators

Chemodetection of common nest predators may be advantageous for nesting birds; however, few studies have examined the ability of songbirds to detect chemical odors from predators. Thus, in 2002, we presented eastern bluebirds Sialia sialis with pairs of nest boxes; one box in the pair was regularly scented with chemical cues from a common nest predator, the black rat snake Elaphe obsoleta , and the other with a neutral cue. In 2004, we again presented bluebirds with pairs of boxes, one scented with chemical cues from a different nest predator, the deer mouse Peromyscus maniculatus , and the other with a neutral scent. Although human females were able to correctly distinguish paper laced with predator cues from paper with neutral cues, bluebirds were as likely to lay eggs in boxes with predator cues as in boxes with neutral cues. While it remains possible that bluebirds may detect scent from potential nest predators, it appears that the presence of these chemical cues does not ultimately influence selection of nest sites.     

Effects of food availability on proceptivity: A test of the reproduction at all costs and metabolic fuels hypotheses

Proceptive behaviours are used by animals to indicate interest in opposite-sex conspecifics. These behaviours can be affected by an individual's nutritional status. Two mutually exclusive hypotheses have been proposed to account for the effects of food availability on reproduction. These are the metabolic fuels hypothesis and the reproduction at all costs hypothesis. It is not known if food availability affects proceptive behaviours such as scent marking, over-marking, and self-grooming. In this study, we tested the hypothesis that food-deprived and nonfood-deprived meadow voles, Microtus pennsylvanicus, differ in the number of scent marks they deposit, the proportion of over-marks they deposit, and the amount of time they spend self-grooming when they encounter the scent marks of opposite-sex conspecifics. We tested this hypothesis by exposing meadow voles that either had continuous access to food or were food-deprived for either 6hours or 24hours to the scent marks of an opposite-sex conspecific. Due to differences in the natural history of male and female meadow voles, we predicted that female voles' behaviour will best be explained by the metabolic fuels hypothesis whereas males' behaviour will best be explained by the reproduction at all costs hypothesis. We found that both male and female voles deprived of food for either 6hours or 24hours spent less time self-grooming compared to nonfood-deprived voles. However, food availability did not affect the scent marking and over-marking behaviour of male and female voles. Differences in the effects of food availability on these proceptive behaviours are discussed within the context of the natural history of meadow voles.     

German Yellowjacket (Vespula germanica) Foragers Use Odors Inside the Nest to Find Carbohydrate Food Sources

Prior work has shown that yellowjacket waSPS remember food odors and use them as cues when foraging. There is also evidence they have mechanisms to recruit nest mates to highly rewarding food sources, as naïve individuals are more likely to go to food sources with scents similar to those visited by nest mates. We asked whether recruitment requires behavioral stimulation by returning foragers, as in honey bees, or if sampling the food source inside the nest is sufficient. We tested this by eliminating the behavior of returning foragers by inserting a scented sugar solution directly into a Vespula germanica nest. Exiting foragers were given a choice of the test scent and a control scent. WaSPS were more likely to choose the test scent. We conclude that behavioral interactions with returning foragers are not necessary to stimulate nest mates to associate an odor with a food source and search for a resource bearing that odor, and that experience with the scented reward inside the nest is sufficient to achieve this result.     

Floral visitors and ant scent marks: noticed but not used?

1. Bee behaviour when visiting flowers is mediated by diverse chemical cues and signals, from the flower itself and from previous visitors to the flower. Flowers recently visited by bees and hoverflies may be rejected for a period of time by subsequent bee visitors. 2. Nectar‐thieving ants also commonly visit flowers and could potentially influence the foraging decisions of bees, through the detection of ant trail pheromones or footprint hydrocarbons. 3. Here we demonstrate that, while naÏve bumblebees in laboratory trials are not inherently repelled by ant scent marks, they can learn to use them as informative signals while foraging on artificial flowers. 4. To test for similar activity in the wild, visitor behaviours at the flowers of Digitalis purpurea Linnaeus, Bupleurum fruticosum Linnaeus, and Brassica juncea (Linnaeus) Czernajew were compared between flowers that had been in contact with ants and those that had not. No differences were found between the two treatments. 5. The use of chemical foraging cues by bees would appear to be strongly dependent on previous experience and in the context of these plant species bees did not associate ant scent mark cues with foraging costs.     

Olfactory learning in the stingless bee Tetragonisca angustula (Hymenoptera, Apidae, Meliponini)

Tetragonisca angustula stingless bees are considered as solitary foragers that lack specific communication strategies. In their orientation towards a food source, these social bees use chemical cues left by co-specifics and the information obtained in previous foraging trips by the association of visual stimuli with the food reward. Here, we investigated their ability to learn the association between odors and reward (sugar solution) and the effect on learning of previous encounters with scented food either inside the hive or during foraging. During food choice experiments, when the odor associated with the food was encountered at the feeding site, the bees’ choice is biased to the same odor afterwards. The same was not the case when scented food was placed inside the nest. We also performed a differential olfactory conditioning of proboscis extension response with this species for the first time. Inexperienced bees did not show significant discrimination levels. However, when they had had already interacted with scented food inside the hive, they were able to learn the association with a specific odor. Possible olfactory information circulation inside the hive and its use in their foraging strategies is discussed.     

Complex memories in honeybees: can there be more than two?

Foraging honeybees are likely to learn visual and chemical cues associated with many different food sources. Here, we explore how many such sources can be memorized and recalled. Marked bees were trained to visit two (or three) sugar feeders, each placed at a different outdoor location and carrying a different scent. We then tested the ability of the bees to recall these locations and fly to them, when the training scents were blown into the hive, and the scents and food at the feeders were removed. When trained on two feeder locations, each associated with a different scent, the bees could correctly recall the location associated with each scent. However, this ability broke down when the number of scents and feeder locations was increased to three. Performance was partially restored when each of the three training feeders was endowed with an additional cue, namely, a distinct colour. Our results suggest that bees can recall a maximum of two locations when each is associated with a different scent. However, this number can be increased if the scent cues are augmented by visual cues. These findings have implications for the ways in which associations are established and laid down in honeybee memory.     

Bumble bee olfactory information flow and contact-based foraging activation

Nestmate foraging activation and interspecific variation in foraging activation is poorly understood in bumble bees, as compared to honey bees and stingless bees. We therefore investigated olfactory information flow and foraging activation in the New World bumble bee species, Bombus impatiens. We (1) tested the ability of foragers to associate forager-deposited odor marks with rewarding food, (2) determined whether potential foragers will seek out the food odor brought back by a successful forager, and (3) examined the role of intranidal tactile contacts in foraging activation. Bees learned to associate forager-deposited odor marks with rewarding food. They were significantly more attracted to an empty previously rewarding feeder presented at a random position within an array of eight previously non-rewarding feeders. However, foragers did not exhibit overall odor specificity for short-term, daily floral shifts. For two out of three tested scents, activated foragers did not significantly prefer the feeder providing the same scent as that brought back by a successful forager. Finally, bees contacted by the successful forager inside the nest were significantly more likely to leave the nest to forage (38.6% increase in attempts to feed from empty feeders) than were non-contacted bees. This is the first demonstration that tactile contact, a hypothesized evolutionary basal communication mechanism in the social corbiculate bees, is involved in bumble bee foraging activation. Received 4 September 2007; revised 30 May 2008; accepted 15 July 2008.     

Stingless bees (<Emphasis Type="Italic">Scaptotrigona pectoralis</Emphasis>) learn foreign trail pheromones and use them to find food

Foragers of several species of stingless bees (Hymenoptera, Apidae and Meliponini) deposit pheromone marks in the vegetation to guide nestmates to new food sources. These pheromones are produced in the labial glands and are nest and species specific. Thus, an important question is how recruited foragers recognize their nestmates’ pheromone in the field. We tested whether naïve workers learn a specific trail pheromone composition while being recruited by nestmates inside the hive in the species Scaptotrigona pectoralis. We installed artificial scent trails branching off from trails deposited by recruiting foragers and registered whether newly recruited bees follow these trails. The artificial trails were baited with trail pheromones of workers collected from foreign S. pectoralis colonies. When the same foreign trail pheromone was presented inside the experimental hives while recruitment took place a significant higher number of bees followed the artificial trails than in experiments without intranidal presentation. Our results demonstrate that recruits of S. pectoralis can learn the composition of specific trail pheromone bouquets inside the nest and subsequently follow this pheromone in the field. We, therefore, suggest that trail pheromone recognition in S. pectoralis is based on a flexible learning process rather than being a genetically fixed behaviour.     

Recognition of scent marks in solitary bees to avoid previously visited flowers

Knowing how floral visitors forage efficiently among flowers is important to understanding plant-pollinator interactions. When bees search for rewarding flowers, they use several visual cues to detect the available floral resources. In addition to these cues, bees can recognize scent marks, which are olfactory cues left on flowers foraged by previous visitors. This behavior is well known in social bees, such as honeybees and bumblebees. Although solitary bees do not need to give information about which flowers were foraged to conspecifics, several pieces of evidence have indicated the use of scent marks. However, it is unknown whether the behavior is widely used in many different bee species. We investigated whether four different solitary bees, Colletes patellatus (Colletidae), Andrena prostomias (Andrenidae), Osmia orientalis (Megachilidae), and Tetralonia mitsukurii (Apidae), can recognize flowers that have been foraged previously by visitors within 3 min. All four bees showed rejection responses to flowers foraged by conspecifics. However, our results showed that responses to foraged flowers varied among bee species. The tendency of A. prostomias and T. mitsukurii to reject the foraged flowers was pronounced, while in C. patellatus and O. orientalis it was weak. In both A. prostomias and T. mitsukurii, the rejection rate of flowers foraged by conspecifics decreased as the time lag after the last visit increased. Both bees visited the flowers from which pollen or nectar had been artificially removed. We suggest that A. prostomias and T. mitsukurii would recognize scent marks left by previous visitors, while the other two bees would not recognize them so strongly. It is likely that the decision to use scent marks is dependent either on the richness of resources or on the complexity of floral structure.     

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.     

Pheromone paths attached to the substrate in meliponine bees: helpful but not obligatory for recruitment success

In contrast to marking of the location of resources or sexual partners using single-spot pheromone sources, pheromone paths attached to the substrate and assisting orientation are rarely found among flying organisms. However, they do exist in meliponine bees (Apidae, Apinae, Meliponini), commonly known as stingless bees, which represent a group of important pollinators in tropical forests. Worker bees of several Neotropical meliponine species, especially in the genus Scaptotrigona Moure 1942, deposit pheromone paths on substrates between highly profitable resources and their nest. In contrast to past results and claims, we find that these pheromone paths are not an indispensable condition for successful recruitment but rather a means to increase the success of recruiters in persuading their nestmates to forage food at a particular location. Our results are relevant to a speciation theory in scent path-laying meliponine bees, such as Scaptotrigona. In addition, the finding that pheromone path-laying bees are able to recruit to food locations even across barriers such as large bodies of water affects tropical pollination ecology and theories on the evolution of resource communication in insect societies with a flying worker caste.     

Female meadow voles,Microtus pennsylvanicus,do not alter their over-marking in response to female conspecifics that were food deprived

Many terrestrial mammals will deposit scent marks and over-marks, the latter being the overlapping scent marks of two conspecifics. Studies have shown that male rodents that are exposed to the overlapping scent marks of two female conspecifics later spend more time investigating the mark of the top-scent female than that of the bottom-scent female. This suggests that over-marking is a form of competition and that the top-scent female is more likely than the bottom-scent female to be chosen as a potential mate. Thus, females should over-mark the scents of neighboring females at a rate that will maximize their chances of attracting males. However, meadow voles live in areas that may contain patchy food availability and residents may differ in their nutritional status. Females in a better nutritional state may be more likely than those in poorer nutritional states to indicate their presence in an area, signal possession of a territory, and to attract mates. Thus, we tested the prediction that female meadow voles, Microtus pennsylvanicus, that were not food deprived would deposit more over-marks than female voles that were food deprived for 6 h. Food-deprived female voles and female voles that had continuous access to food deposited a similar number of scent marks and used a similar proportion of those marks as over-marks when they encountered the scent marks of female conspecifics. These findings suggest that the nutritional status of female voles does not affect their ability to signal their presence in an area marked by a female conspecific.     

Ants use pheromone markings in emigrations to move closer to food-rich areas

Nest site quality can affect survival and reproduction, and thus many animals have evolved behaviors which facilitate nest site assessment and selection. Ants of the genus Temnothorax have been shown to include an array of nest site attributes when choosing such a site. Here, we show that they also include traits of the habitat surrounding nest sites. In particular, we found that during emigration, ants preferred to move to nests located close to a previously explored food-rich area. We also determined that scent markings played a role in this choice and that scouts and transporting ants may have tracked scent marks laid in foraging, and this behavior could have biased emigration toward nests located near previously foraged areas. These results indicate that pheromones play a bigger role in Temnothorax foraging and decision making in emigration than previously thought. Overall, this work provides new insights into the mechanisms involved in habitat selection in ants and contributes to our understanding of collective behavior in social insects in general.     

Meadow voles and prairie voles differ in the percentage of conspecific marks they over-mark

Many terrestrial mammals scent mark in areas containing the scent marks of conspecifics, and thus, may deposit their own scent marks on top of those that were deposited previously by conspecifics. This phenomenon, known as over-marking appears to play a role in same-sex competition or mate attraction. The present study determines whether meadow and prairie voles avoid over-marking the scent marks of conspecifics, target the scent marks of conspecifics and over-mark them, or randomly over-mark the scent marks of conspecifics. The data show that meadow and prairie voles adjust the number and location of scent marks that they deposit in areas marked previously by particular conspecifics. Male and female meadow and prairie voles target the scent marks of opposite-sex conspecifics and over-mark them. Female meadow and prairie voles also target the scent marks of female conspecifics. In contrast, male meadow and prairie voles over-mark the scent marks of male conspecifics in a random manner. By differentially over-marking the scent marks of conspecifics, voles may be able to communicate particular information to a variety of conspecifics.     

Individually recognizable scent marks on flowers made by a solitary bee

The marking of flowers with ephemeral scent is an underappreciated but vital element in the foraging behaviour of social bees. Using observational and experimental data, we tested whether a solitary bee (female Anthophora plumipes) uses scent marking while foraging on flowers of Cerinthe major in Portugal. Females used scent marks with at least two components that differed in their volatility and, furthermore, recognized the marks of different individuals. A very short-term component (<3 min) was attractive, resulting in the observed high level of immediate revisits: this component appeared to be adjusted according to the foraging needs of the moment. A longer-term component (<30 min) was initially repellent and matched the rate of nectar renewal; it, or the response to it, also appeared to be adjusted to the perceived level of nectar reward. There may be even longer-term effects associated with the specific foraging areas of individual bees. Observed differences in the way in which individuals responded to scent marks indicate that they may play a role as part of a dominance or exclusion mechanism among females. Copyright 2001 The Association for the Study of Animal Behaviour.     

Female marmoset monkeys (Callithrix jacchus) can be identified from the chemical composition of their scent marks.

The present study analyzed 42 organic solvent extracts of scent mark pools from five dominant female common marmosets by gas chromatography (GC) and combined GC and mass spectrometry. We determined whether there were qualitative or quantitative differences between the chemical composition of scent marks from individual females. Gas chromatography and mass spectral analysis detected the same 162 chemicals in 86% (36/42) of scent mark pools from five dominant females. This near identical chemical composition of scent marks suggested there were few, if any, qualitative differences between the chemical composition of scent marks from individual females. Instead, quantitative differences in scent may provide the key factor distinguishing individual females. Using the relative concentration of highly volatile chemicals detected by GC in scent marks, linear discriminant analysis classified scent mark pools to their correct donor approximately 91% of the time. Such highly reliable statistical matching of scent to donor suggested that each individual female common marmoset has a unique ratio of highly volatile chemicals in their scent marks which may permit individual identification of females from odors in their scent alone.     

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.     

Orientation and Communication in the Neotropical Ant Odontomachus bauri Emery (Hymenoptera,Formicidae, Ponerinae)

The Neotropical species Odontomachus bauri employs canopy orientation during foraging and homing. An artificial canopy pattern above the ants is much more effective as an orientation cue than horizontal landmarks or chemical marks. However, both horizontal visual cues and chemical marks on the ground can serve in localizing the nest entrance. Successful O. bauri foragers recruit nestmates to leave the nest and search for food. However, the recruitment signals do not contain directional information. Antennation bouts and pheromones from the pygidial gland most likely serve as stimulating recruitment signals. Secretions from the mandibular and poison gland elicit alarm and attack behavior.     

Contextual modulation of visuomotor associations in bumble-bees (Bombus terrestris)

We performed the following experiment to investigate whether contextual cues can prevent interference during the acquisition of potentially competing visuomotor associations. In the bumble-bee (Bombus terrestris). Bees discriminated between horizontally and vertically orientated gratings of black and white stripes in order to reach a feeder and between different diagonally orientated gratings to gain access to their nest. Once bees were well trained on these two tasks, the discrimination task at the nest was changed so that bees had to distinguish between horizontal and vertical gratings at both sites. Whereas they still approached the horizontal grating to reach food, they now had to approach a vertical grating in order to return to their nest. The new task was learnt rapidly. Errors at the feeder did not increase during or after the acquisition of this potentially competing behaviour at the nest entrance. For a brief period during acquisition, bees showed some hesitation at the feeder and hovered for slightly longer before choosing between horizontals and verticals. After acquisition was complete, bees showed a slight increase in their preference for verticals over a more ambiguous stimulus of an array of dots. These findings are consistent with the hypothesis that different contextual signals are associated with approaching the nest or approaching the feeder, and that these contextual signals facilitate learnt associations between orientation detectors and motor commands.     

Intraspecific Variation of Floral Scent Chemistry in Magnolia kobus DC. (Magnoliaceae)

Magnolia kobus was examined at 32 sites in Japan (109 female-stage flowers from 52 plants) by GC-MS. Major chemical compounds (a total of 36 chemicals) emitted from the flowers were: linalool (and its oxides), limonene, cis- and trans-β-Ocimene, benzaldehyde, benzyl alcohol, benzyl cyanide, and 2-aminobenzaldehyde. Linalool and its oxides were the most abundant components of floral scents in 21 individuals. The rate at which chemical volatiles were emitted ranged from 0.002 to 0.929 μg/flower/hour (average 0.211). High quantitative and qualitative variation in floral scent chemistry among individuals was found throughout the range of M. kobus, especially in central Honshu. The high variability in floral scent chemistry may be due to the importance of visual cues in the reproductive biology of M. kobus which flowers in early spring, resulting in decreased selection for specific floral scent profiles. Alternatively, different scent compounds or chemical profiles may be equally effective in attracting pollinators. Received 25 June 2001/ Accepted in revised form 25 August 2001