Olfactory response of megachilid bees, Osmia lignaria, Megachile rotundata, and M. pugnata, to individual cues from old nest cavities

The megachilid bees Osmia lignaria Say, Megachile rotundata (F.), and M. pugnata Say were tested for attraction to various components associated with their old nest cavities, or chemical extracts of these components, using a Y-tube olfactory response bioassay. Female bees of these species are known to nest in or near old nest cavities, implying that remnant nest components are important cues for bees looking for nest cavities. Significant results show that female bees were attracted to components that may provide species-specific cues or indicate conspecific nesting activity. Specifically, O. lignaria females showed attraction only to the female cocoon. M. rotundata females were attracted to intact nest cells, the fecal material on the outside of a cocoon, leaf pieces used as nest cell lining, and the extract of leaf pieces. M. pugnata females were attracted to the whole nest cell, the paper straw nesting material with attached cocoon, and feces.     

Individual scent marking of the nest entrance as a mechanism for nest recognition inXylocopa pubescens (Hymenoptera: Anthophoridae)

The mechanism by which female Xylocopa pubescenslocate their nest in a nesting aggregation was investigated. The bees were induced to nest in canes to which uniform nest entrances were attached. The results of nest displacement experiments revealed that the bees use visual cues for proximate orientation,but at very close range they also use olfactory cues. This conclusion was corroborated by the results of experiments in which the nest entrances were either removed or exchanged for alien nest entrances. Moreover, habituation experiments strongly indicated that the bees impart their individual marking at the nest entrance and that they can learn and memorize the individual odors of the neighboring bees.     

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.     

The role of nest‐site selection and cereal production in differential nest predation in Common Quail Coturnix coturnix and hybrid quail C. coturnix × C. japonica

In Europe, farm‐reared quail used for restocking purposes are often hybrids between the Common Quail Coturnix coturnix and the Japanese Quail Coturnix japonica. These hybrids interbreed with wild Common Quail populations and suffer higher rates of nest predation, which would suggest that the two quail types have different nest‐site selection patterns. We monitored 103 radiotagged nesting females (77 wild Common Quail over 18 breeding seasons and 26 hybrid quail over six breeding seasons) and analysed their nest placement behaviour. Our results did not provide any evidence that the quail types showed different nest‐site selection trends, as they all preferred to nest near cereal field margins. The higher nest predation rate experienced by hybrids might therefore be explained by less effective anti‐predator behaviour among hybrid females during incubation. As no edge effect on nest predation was observed, the preference for nesting close to cereal margins could be due to the greater food resources of field margins. Moreover, Common Quail nest predation decreased with a rise in barley straw production, supporting evidence that nest cover plays an important role in the prevention of nest predation in this species. No such relationship was found for hybrid quail, but this result is not conclusive, as the data on barley straw production for hybrid quail covered a much smaller range of interannual variability. Barley straw production in Catalonia decreased significantly between 1992 and 2012, possibly as a result of a change in the barley varieties used. This practice could entail a conservation threat for ground‐nesting bird species.     

Buildings promote higher incubation temperatures and reduce nest attentiveness in a Neotropical thrush

Incubation is an energetically costly parental task of breeding birds. Incubating parents respond to environmental variation and nest‐site features to adjust the balance between the time spent incubating (i.e. nest attentiveness) and foraging to supply their own needs. Non‐natural nesting substrates such as human buildings impose new environmental contexts that may affect time allocation of incubating birds but this topic remains little studied. Here, we tested whether nesting substrate type (buildings vs. trees) affects the temperature inside the incubation chamber (hereafter ‘nest temperature’) in the Pale‐breasted Thrush Turdus leucomelas, either during ‘day’ (with incubation recesses) or ‘night’ periods (representing uninterrupted female presence at the nest). We also tested whether nesting substrate type affects the incubation time budget using air temperature and the day of the incubation cycle as covariates. Nest temperature, when controlled for microhabitat temperature, was higher at night and in nests in buildings but did not differ between daytime and night for nests in buildings, indicating that buildings partially compensate for incubation recesses by females with regard to nest temperature stability. Females from nests placed in buildings exhibited lower nest attentiveness (the overall percentage of time spent incubating) and had longer bouts off the nest. Higher air temperatures were significantly correlated with shorter bouts on the nest and longer bouts off the nest, but without affecting nest attentiveness. We suggest that the longer bouts off the nest taken by females of nests in buildings is a consequence of higher nest temperatures promoted by man‐made structures around these nests. Use of buildings as nesting substrate may therefore increase parental fitness due to a relaxed incubation budget, and potentially drive the evolution of incubation behaviour in certain urban bird populations.     

Are olfactory cues involved in nest recognition in two social species of estrildid finches?

Reliably recognizing their own nest provides parents with a necessary skill to invest time and resources efficiently in raising their offspring and thereby maximising their own reproductive success. Studies investigating nest recognition in adult birds have focused mainly on visual cues of the nest or the nest site and acoustic cues of the nestlings. To determine whether adult songbirds also use olfaction for nest recognition, we investigated the use of olfactory nest cues for two estrildid finch species, zebra finches (Taeniopygia guttata) and Bengalese finches (Lonchura striata var. domestica) during the nestling and fledgling phase of their offspring. We found similar behavioural responses to nest odours in both songbird species. Females preferred the odour of their own nest over a control and avoided the foreign conspecific nest scent over a control during the nestling phase of their offspring, but when given the own odour and the foreign conspecific odour simultaneously we did not find a preference for the own nest odour. Males of both species did not show any preferences at all. The behavioural reaction to any nest odour decreased after fledging of the offspring. Our results show that only females show a behavioural response to olfactory nest cues, indicating that the use of olfactory cues for nest recognition seems to be sex-specific and dependent on the developmental stage of the offspring. Although estrildid finches are known to use visual and acoustic cues for nest recognition, the similar behavioural pattern of both species indicates that at least females gain additional information by olfactory nest cues during the nestling phase of their offspring. Thus olfactory cues might be important in general, even in situations in which visual and acoustic cues are known to be sufficient.     

Kea Nestor notabilis mothers produce nest-specific calls with low amplitude and high entropy

Vocal behaviour of nesting altricial birds is subject to selection pressure from several sources. Offspring beg to attract parents’ attention, thus increasing the chances of being fed, but also increasing the chances of being detected by predators. Research on passerines has shown that parents may reduce the risk of nest predation by alarm calling to warn nestlings to be quiet, and by producing food calls which solicit begging when parents are present to defend the nestlings. Both nestlings and parents may reduce the risk of predator detection by producing calls of low amplitude and high entropy which are acoustically difficult to locate. Although extensive research has been undertaken on nesting passerine vocalizations, little is known about parrots in this regard, and studies are needed to determine whether parrots show similar adaptations. We investigated the calling behaviour of Kea Nestor notabilis mothers during the nesting period to determine whether maternal vocalizations were adapted in a way that could increase the chance of brood success. A microphone was installed inside the nest to record calls produced both inside the nest and in the direct vicinity. Our prediction was that calls outside the nest would be easy to locate and could function as alarm calls to alert conspecifics or distract the predator, whereas calls inside the nest would be difficult to locate and could serve to communicate with nestlings without alerting predators. Our results accorded with these predictions. Calls produced outside the nest were loud and tonal, and corresponded to previously described Kea alarm calls. Calls produced inside the nest, however, were high-entropy and low-amplitude calls, and formed a distinct structural category. We thus provide the first evidence that a parrot species has a vocal category for communication inside the nest, and that calls within this category are structured in a way that could reduce the risk of nest predation.     

How Do Infanticidal Male Bank Voles (Myodes glareolus) Find the Nest with Pups?

Infanticide, the killing of conspecific young, occurs in most mammal species, like in our study species, the bank vole (Myodes glareolus). Infanticide by adult males is regarded as a strong factor affecting recruitment of young into population. It is considered as an adaptive behaviour, which may increase male fitness via resource gain or an increased access to mates. When an intruder is approaching the nest, the mother should not be present, as her nest guarding is very aggressive and successful. Pups use ultrasonic vocalisation to call their mother when mother leaves nest for foraging but it is not know which cues do infanticidal males use to find the nest with vulnerable pups to commit infanticide? We studied whether the pups' sounds or the olfactory cues of the nest guide the males of known infanticidal behavioural trait towards the nest with vulnerable pups. Four nest boxes in a large indoor arena offered different nesting cues: nest odour, pup vocalisation, both odour and sound or control with no cue. The result showed that infanticidal males were more active in their searching behaviour than non‐infanticidal males and seemed to target the nest providing only acoustic cues. Four of the males, all infanticidal, intruded the nest box. Infanticidal males seem to actively search for nests with vulnerable pups by eavesdropping pup begging calls for absent mother. However, under natural conditions, mother presence and aggressive nest protection may be an effective counter strategy against strange male infanticide. When trapping study voles from the wild, we monitored occurrence of male infanticide across the breeding season from early to late summer. Proportion of infanticidal males was between 25 and 29% of all males tested along the breeding season. Our results suggest that male infanticide seems to cause a stable threat for pup mortality in increasing breeding season density.     

Nest height is affected by lamppost lighting proximity in addition to nestbox size in urban great tits

Both natural and artificial light have proximate influences on many aspects of avian biology, physiology and behaviour. To date artificial light at night is mostly considered as being a nuisance disrupting for instance sleep and reproduction of diurnal species. Here, we investigate if lamppost night lighting affects cavity‐nesting bird species inside their breeding cavity. Nest height in secondary cavity‐nesting species is the result of trade‐offs between several selective forces. Predation is the prevailing force leading birds to build thin nests to increase the distance towards the entrance hole. A thin nest may also limit artificial light exposure at night. Yet, a minimum level of daylight inside nesting cavities is necessary for adequate visual communication and/or offspring development. Against this background, we hypothesised that avian nest‐building behaviour varies in response to a change in night lighting. We monitored nest height of urban great tits Parus major during six years and found that it varied with artificial light proximity. The birds built thinner nests inside nestboxes of various sizes in response to increasing lamppost night light availability at the nest. In large nestboxes, the nests were also thinner when a lamppost was present in the territory. Whether this relationship between artificial night lighting and nest height reflects a positive or negative effect of urbanisation is discussed in the light of recent experimental studies conducted in rural populations by other research groups.     

Spatial Nest‐Settlement Decisions in Digger Wasps: Conspecifics Matter more than Heterospecifics and Previous Experience

Research into the driving forces behind spatial arrangement of wasp nests has considered abiotic environmental factors, but seldom investigated attraction or repulsion towards conspecifics or heterospecifics. Solitary female digger wasps (Hymenoptera) often nest in dense aggregations, making these insects good models to study this topic. Here, we analysed the nesting patterns in an area shared by three species of the genus Bembix, in a novel study to discover whether female wasps are attracted to or repulsed by conspecific nests, heterospecific nests or their own previously established nests when choosing nest‐digging locations. Early in the season, each species showed a clumping pattern of nests, but later in the season, a random distribution of nests was more common, suggesting an early conspecific attraction. Such behaviour was confirmed by the fact that females started building their nests more frequently where other females of their species were simultaneously digging. The distances between subsequent nests dug by individual females were shorter than those obtained by random simulations. However, this pattern seemed to depend on the tendency to dig close to conspecifics rather than remain in the vicinity of previous nests, suggesting that females' experience matters to future decisions only on a large scale. Nesting patches within nest aggregations largely overlapped between species, but the nests of each species were generally not closer to heterospecific nests than expected by chance, suggesting that females are neither repulsed by, nor attracted to, congenerics within nest aggregations. A role of the spatial distribution of natural enemies on the observed nesting patterns seemed unlikely. Bembix digger wasp nest aggregations seem thus to be primarily the result of female–female attraction during nest‐settlement decisions, in accordance with the ‘copying’ mechanisms suggested for nesting vertebrates.     

Does variation in female body size affect nesting success in Dawson’s burrowing bee,Amegilla dawsoni (Apidae: Anthophorini)?

Abstract.  1. Females of Dawson's burrowing bees vary in body weight over a twofold range. Despite the potential for differences in body weight to affect several aspects of the competition among nesting females, no clear advantages were documented for larger females.
2. Nesting females were not consistently larger than emerging females, nor was there a consistent relationship between body size and the weight of pollen and nectar carried to the nest on provisioning trips. At one nesting location, larger females did not produce larger pre-pupal offspring nor did they produce offspring at a faster rate than their smaller nesting companions.
3. In addition, large body size was not associated with greater success in nest defence despite the fact that nesting females regularly encountered intruders in their burrows. Residency, not body size, determined the outcome of almost all contests for control of a nest burrow. The absence of a large body size effect here appears to stem from an intruder strategy designed to enable nest-searching females to acquire burrows that had been abandoned through death or dispersal of their original owners rather than securing existing nests through an aggressive takeover strategy.
4. Thus, although large body size conveys significant fitness advantages to males, this attribute does not promote female success in either provisioning or defending their nests.     

Dispersed versus gregarious nesting strategies in the mason bee Chalicodoma siculum

Chalicodoma siculum (Rossi) is a mason bee that usually nests in dispersed sites, constructing spherical mud nests on twigs or hemispherical nests on hard surfaces. Under favourable conditions it may nest in aggregations of several hundred bees. At the nesting aggregation, nest usurpation often occurs, either by the laying of eggs in finished but unsealed cells, by opening a sealed cell and replacing its egg, or by taking over an unfinished cell. Mutual construction and provisioning of a single cell by several females was also often observed. Because participant bees display extreme aggression in prolonged disputes over cell ownership, these interactions can be regarded as competitive rather than cooperative. Moreover, since the total investment in such cells is not different from that in cells located in dispersed sites, the female that actually manages to lay an egg gains by participating in a cell under construction.
We discuss the advantages of nesting in aggregations vs. the risks of being usurped, as selective forces underlying nesting behaviour of bees.     

Nesting aggregation as a predictor of brood parasitism in mason bees (Osmia spp.)

1. Parasitism can be an important source of mortality for insect populations; however, we know little about the factors influencing vulnerability of wild bees to parasites. Mason bees (genus Osmia; Hymenoptera: Megachilidae) are important pollinators of crops and wild plants and are vulnerable to attack by brood parasites. High nest densities may increase rates of brood parasitism by attracting disproportionate numbers of parasites. 2. Three years of field observations from multiple sites were analysed to assess whether mason bee brood parasitism increased with host density. Mason bees were allowed to nest in artificial nesting blocks and establish natural variation in nesting density. Nest cells constructed by bees were checked for the presence of parasite eggs. 3. Parasitism of nest cells strongly increased with the number of actively nesting bees at a nesting block. Mason bees showed no preference for nesting in blocks that were occupied or unoccupied by other mason bees. Parasitism also increased with the number of days a nest was provisioned and decreased over the course of the season. Nest cells constructed last in a nest were significantly more parasitised than inner cells, despite being sealed against invasions. 4. These findings show positively density‐dependent parasitism in mason bees. They also suggest that bees terminate parasitised nests, causing parasitised cells to become outermost nest cells – a behaviour that may represent a defence against parasites. Our results have implications for the management of mason bees as agricultural pollinators, as cultivating them at high densities could reduce offspring survival.     

Kin grouping is insufficient to explain the inclusive fitness gains of conspecific brood parasitism in the common eider

Conspecific brood parasitism allows females to exploit other females' nests and enhance their reproductive output. Here, we test a recent theoretical model of how host females gain inclusive fitness from brood parasitism. High levels of relatedness between host and parasitizer can be maintained either by: (a) kin recognizing and parasitizing each other as a form of cooperative breeding or (b) natal philopatry and nest site fidelity facilitating the formation of kin groups, thereby increasing the probability of parasitism between relatives nesting in close proximity. To address these two hypotheses we genotyped feathers and hatch membranes of common eiders (Somateria mollissima) from western Hudson Bay, Canada, using a noninvasive sampling methodology. We found that most instances of brood parasitism do result in inclusive fitness gains. Furthermore, females with failed nests moved an average of 492 m from their previous year's nest site, while successful females only moved an average of 13 m. Therefore, we observed host–parasite relatedness can occur at levels higher than would be expected by chance even in the absence of kin grouping, suggesting that closely related females nesting near one another is not essential to maintain high host–parasitizer relatedness. In addition, kin grouping is only a transient phenomenon that cannot occur every year due to the propensity for females of failed nests to nest farther away from their nest site in subsequent years than females with successful nests, which provides support for kin recognition as a more likely mechanism to maintain high host–parasitizer relatedness over time.     

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.     

Climate change and nesting behaviour in vertebrates: a review of the ecological threats and potential for adaptive responses

Nest building is a taxonomically widespread and diverse trait that allows animals to alter local environments to create optimal conditions for offspring development. However, there is growing evidence that climate change is adversely affecting nest‐building in animals directly, for example via sea‐level rises that flood nests, reduced availability of building materials, and suboptimal sex allocation in species exhibiting temperature‐dependent sex determination. Climate change is also affecting nesting species indirectly, via range shifts into suboptimal nesting areas, reduced quality of nest‐building environments, and changes in interactions with nest predators and parasites. The ability of animals to adapt to sustained and rapid environmental change is crucial for the long‐term persistence of many species. Many animals are known to be capable of adjusting nesting behaviour adaptively across environmental gradients and in line with seasonal changes, and this existing plasticity potentially facilitates adaptation to anthropogenic climate change. However, whilst alterations in nesting phenology, site selection and design may facilitate short‐term adaptations, the ability of nest‐building animals to adapt over longer timescales is likely to be influenced by the heritable basis of such behaviour. We urgently need to understand how the behaviour and ecology of nest‐building in animals is affected by climate change, and particularly how altered patterns of nesting behaviour affect individual fitness and population persistence. We begin our review by summarising how predictable variation in environmental conditions influences nest‐building animals, before highlighting the ecological threats facing nest‐building animals experiencing anthropogenic climate change and examining the potential for changes in nest location and/or design to provide adaptive short‐ and long‐term responses to changing environmental conditions. We end by identifying areas that we believe warrant the most urgent attention for further research.     

Predation Cues in Solitary bee Nests

Predation at the nesting site can significantly affect solitary bees’ reproductive success. We tested female red mason bees’ (Osmia bicornis L.) acceptance of potential nesting sites, some of which were marked with cues coming from predated conspecifics (crushed bees) or from a predator itself (rodent excreta). In our experiment, females did not avoid nests marked with either of the two predator cues. We suggest that bee females do not recognize these two cues as risky. Alternatively, costs of abandoning natal aggregation might be too high compared with any perceived predation risk of staying. Moreover, the presence of crushed bees can provide positive information about the presence of conspecifics and, possibly, information about a nesting aggregation that may be preferred by bees when choosing a nesting site.     

Evolution of nesting behaviour and kleptoparasitism in a selected group of osmiine bees (Hymenoptera: Megachilidae)

The construction of nests to rear offspring is restricted to vertebrates and few insect taxa, such as termites, wasps, and bees. Among bees, species of the family Megachilidae are characterized by a particularly high diversity in nest construction behaviour. Many megachilid bees nest in excavated burrows in the ground, others place their brood cells in a variety of above‐ground cavities or attach them to the surface of a substrate, and yet others have adopted a kleptoparasitic habit. Evolutionary transitions between the different nesting sites and between conventional nesting and kleptoparasitism in bees are poorly understood. In the present study, we traced the evolution of nesting site selection and kleptoparasitism in the Annosmia–Hoplitis group (Osmiini), which displays an exceptionally high diversity in nesting behaviour. We found that the evolution of nesting behaviour proceeded unidirectionally from nesting in excavated burrows in the ground to nesting in rock depressions and cavities, followed by the colonization of snail shells and insect borings in dead wood or hollow stems. Kleptoparasitism evolved once and the kleptoparasitic species have derived from the same lineage as their hosts. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 349–360.     

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.