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.     

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.     

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.     

Conditional female strategies influence hatching success in a communally nesting iguana

The decision of females to nest communally has important consequences for reproductive success. While often associated with reduced energetic expenditure, conspecific aggregations also expose females and offspring to conspecific aggression, exploitation, and infanticide. Intrasexual competition pressures are expected to favor the evolution of conditional strategies, which could be based on simple decision rules (i.e., availability of nesting sites and synchronicity with conspecifics) or on a focal individual's condition or status (i.e., body size). Oviparous reptiles that reproduce seasonally and provide limited to no postnatal care provide ideal systems for disentangling social factors that influence different female reproductive tactics from those present in offspring‐rearing environments. In this study, we investigated whether nesting strategies in a West Indian rock iguana, Cyclura nubila caymanensis, vary conditionally with reproductive timing or body size, and evaluated consequences for nesting success. Nesting surveys were conducted on Little Cayman, Cayman Islands, British West Indies for four consecutive years. Use of high‐density nesting sites was increasingly favored up to seasonal nesting activity peaks, after which nesting was generally restricted to low‐density nesting areas. Although larger females were not more likely than smaller females to nest in high‐density areas, larger females nested earlier and gained access to priority oviposition sites. Smaller females constructed nests later in the season, apparently foregoing investment in extended nest defense. Late‐season nests were also constructed at shallower depths and exhibited shorter incubation periods. While nest depth and incubation length had significant effects on reproductive outcomes, so did local nest densities. Higher densities were associated with significant declines in hatching success, with up to 20% of egg‐filled nests experiencing later intrusion by a conspecific. Despite these risks, nests in high‐density areas were significantly more successful than elsewhere due to the benefits of greater chamber depths and longer incubation times. These results imply that communal nest sites convey honest signals of habitat quality, but that gaining and defending priority oviposition sites requires competitive ability.     

Strong selection on mandible and nest features in a carpenter bee that nests in two sympatric host plants

Host plants are used by herbivorous insects as feeding or nesting resources. In wood‐boring insects, host plants features may impose selective forces leading to phenotypic differentiation on traits related to nest construction. Carpenter bees build their nests in dead stems or dry twigs of shrubs and trees; thus, mandibles are essential for the nesting process, and the nest is required for egg laying and offspring survival. We explored the shape and intensity of natural selection on phenotypic variation on three size measures of the bees (intertegular width, wing length, and mandible area) and two nest architecture measures (tunnel length and diameter) on bees using the native species Chusquea quila (Poaceae), and the alloctonous species Rubus ulmifolius (Rosaceae), in central Chile. Our results showed significant and positive linear selection gradients for tunnel length on both hosts, indicating that bees building long nests have more offspring. Bees with broader mandibles show greater fitness on C. quila but not on R. ulmifolius. Considering that C. quila represents a selective force on mandible area, we hypothesized a high adaptive value of this trait, resulting in higher fitness values when nesting on this host, despite its wood is denser and hence more difficult to be bored.     

Early-season colony development of the paper wasp Ropalidia plebeiana (Hymenoptera: Vespidae) in Canberra, Australia

This study presents the first observations of early-season colony development of Ropalidia plebeiana in Canberra, Australia. The growth pattern of three R. plebeiana nests was measured during weekly observations from October 2006 to January 2007 and showed that nests steadily increase in size over the early summer to approximately 50 cells when the nest is newly established and to approximately 170 cells when nests from the previous season are re-used. A first generation of adult females is produced by December, and the bimodality of the curves of egg, larval and pupal numbers indicates that these three developmental stages last approximately 2–3 weeks each. The nesting cycle of R. plebeiana in Canberra commences approximately 2 weeks later than in coastal Australia, the shorter summers in this inland region restricting the length of the active season. R. plebeiana did not form dense nesting aggregations in Canberra as described elsewhere, with only small colonies consisting of a single or just a few nests. Characteristic comb-cutting behavior of the species was observed but this did not result in complete nest division as recorded from coastal populations.     

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.     

Nest site preference and success in a gregarious, ground-nesting bee Dieunomia triangulifera

1. The characteristics of preferred nest sites of the solitary bee Dieunomia triangulifera, which nests gregariously in alluvial soil, were investigated. Nest presence was used as an indication of preference. In those cases where nests were more than a few days old, therefore, this study also investigated nest success. 2. Independent variables tested were soil surface moisture, presence of nesting conspecifics, smooth vs. irregular soil surface, distance to visual landmarks, percentage of vegetation cover, soil compaction, vehicular traffic, soil surface temperature, and light intensity. 3. Bees preferred to nest in moist, compact soil with an irregular surface. More bees nested near visual landmarks and in places with little or no vegetation. They preferred nesting in areas with warmer soil surface temperatures and brighter illumination. The presence of nesting conspecifics did not influence nesting decisions. Vehicular traffic also seemed to have no impact on nest initiation but did seem to have a negative impact on nest success. 4. Although it might seem that ground-nesting bees should not be limited by nesting sites, relatively little of the study area met all their apparent criteria. This might account, in part, for gregarious nesting in this species.     

Influences on the density and dispersion of bumble bee nests (Hymenoptera: Apidae)

To determine some of the influences on nesting densities of bumble bees I examined the dispersion and occupancy of 35 nests in an old field on Amherst Island, Ontario, Canada and 39 nests found by Cumber (1953) in England. Densities up to 1 nest per 200 m² were observed. Overall, Cumber's nests were randomly dispersed, but nests on the ground surface were aggregated. Nests of surface-nesting species were consequently more aggregated than expected at random, whereas surface-nesting and underground-nesting species seldom nested near one another. This distribution of nests probably reflects the nesting habits of the rodents that originally constructed the nests, rather than interactions between bees. On Amherst Island, abnormally wet spring weather apparently hampered normal colony development, particularly of underground-nesting bees based on a low frequency of underground nests, scarcity of workers of two underground-nesting species compared with the relative abundance of spring queens, and delayed appearance of workers of all species relative to the previous year. The nests of two common species were randomly dispersed, but those of a third species were regularly distributed, suggesting that intraspecific interactions may limit nesting density for this species.     

Nest taphonomy of common terns (Sterna hirundo) on Poplar Island,Chesapeake Bay,Maryland

Mesozoic non-avian theropod dinosaurs displayed a diverse range of egg types and clutch forms, suggesting a variety of nesting behaviours, some of which may be shared with birds. More accurate inferences of these behaviours require taphonomic studies of modern nesting sites. Here, we document common tern (Sterna hirundo) nesting sites on Poplar Island in Chesapeake Bay, Maryland. Nests were surveyed on multiple occasions, documenting nest composition, density and distribution, as well as eggshell concentration and orientation. Three colonies yielded 79 tern nests with 193 eggs. Twelve nests hatched, 7 were predated, 30 failed and the fates of 30 remain unknown. Abundant eggshells occurred at the centre of the nests. Concave-up eggshell in or on the nest surface characterised hatched and predated nests, whereas eggshell forced into the subsurface seemed to favour concave-down orientation and may reflect post-hatching extended use of the nest. Eggs buried deep within the nest materials and/or substrate suggest adult abandonment. Quantitative data on eggshell orientation and observations regarding controls on nest distribution, egg predation and incorporation of intact eggs into the substrate at modern nesting sites provide physical guidelines for the improved interpretation of fossil nesting localities.     

Genetic structure at three spatial scales is consistent with limited philopatry in Ricord's Rock Iguanas (Cyclura ricordii)

Cyclura ricordii is an endemic iguana from Hispaniola Island and is threatened on the IUCN Red List. The main threats are predation by introduced mammals, habitat destruction, and hunting pressure. The present study focused on two nesting sites from Pedernales Province in the Dominican Republic. The hypothesis that natal philopatry influences dispersal and nest‐site selection was tested. Monitoring and sampling took place in 2012 and 2013. Polymorphic markers were used to evaluate whether natal philopatry limits dispersal at multiple spatial scales. Ripley's K revealed that nests were significantly clustered at multiple scales, when both nesting sites were considered and within each nesting site. This suggests a patchy, nonrandom distribution of nests within nest sites. Hierarchical AMOVA revealed that nest‐site aggregations did not explain a significant portion of genetic variation within nesting sites. However, a small but positive correlation between geographic and genetic distance was detected using a Mantel's test. Hence, the relationship between geographic distance and genetic distance among hatchlings within nest sites, while detectable, was not strong enough to have a marked effect on fine‐scale genetic structure. Spatial and genetic data combined determined that the nesting sites included nesting females from multiple locations, and the hypothesis of “natal philopatry” was not supported because females nesting in the same cluster were no more closely related to each other than to other females from the same nesting site. These findings imply that nesting aggregations are more likely associated with cryptic habitat variables contributing to optimal nesting conditions.     

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.     

Nest boxes for Cape Parrots Poicephalus robustus in the Hogsback area,Eastern Cape,South Africa

Breeding propensity of tree-cavity nesting bird species are often limited by a shortage of natural nesting sites. Artificial nests can be used to provide alternative nest sites. Cape Parrots Poicephalus robustus are nationally endangered and nest in existing tree-cavities in high-altitude fragmented Afromontane forests in South Africa, assumed to be in short supply due to historic and current logging practices. To increase nest site availability, 179 wooden bird boxes and 28 bee boxes (to ‘pull’ bees) were erected during 2011–2012 in Hogsback, Eastern Cape. In 2016, no bird boxes were occupied by Cape Parrots. A total of 43% were used by other species, 51% were unused and 6% could not be inspected due to tree instability and inaccessibility. Two bird boxes were inspected by two pairs of Cape Parrots, but were never occupied. Occupancy of boxes by birds was not associated with nest, tree or habitat characteristics. However, occupancy of boxes by bees was associated with habitat type and tree species. Future conservation efforts will include locating natural Cape Parrot nesting sites and reforestation efforts to ensure the long-term availability of natural nesting sites.     

‘Bee Hotels’ as Tools for Native Pollinator Conservation: A Premature Verdict?

Society is increasingly concerned with declining wild bee populations. Although most bees nest in the ground, considerable effort has centered on installing ‘bee hotels’—also known as nest boxes or trap nests—which artificially aggregate nest sites of above ground nesting bees. Campaigns to ‘save the bees’ often promote these devices despite the absence of data indicating they have a positive effect. From a survey of almost 600 bee hotels set up over a period of three years in Toronto, Canada, introduced bees nested at 32.9% of sites and represented 24.6% of more than 27,000 total bees and wasps recorded (47.1% of all bees recorded). Native bees were parasitized more than introduced bees and females of introduced bee species provisioned nests with significantly more female larva each year. Native wasps were significantly more abundant than both native and introduced bees and occupied almost 3/4 of all bee hotels each year; further, introduced wasps were the only group to significantly increase in relative abundance year over year. More research is needed to elucidate the potential pitfalls and benefits of using bee hotels in the conservation and population dynamics of wild native bees.     

Communal egg-laying behaviour and the consequences of egg aggregation in the brown anole (Anolis sagrei)

Communal nesting is a behaviour that involves multiple females laying eggs in the same nest or nesting site. This behaviour may be a consequence of a shortage of preferable nest sites (constraint hypothesis) or an adaptation generated by fitness benefits associated with egg aggregation (adaptive hypothesis). Experimental tests of these hypotheses require information about maternal nest site choice and its fitness consequences. To address these, we studied a lizard (brown anole; Anolis sagrei) that produces single-egg clutches, but often aggregates eggs in nest sites. In a lab study, females were given the option of nesting in (a) soil previously used as nest substrate vs. fresh soil and (b) soil with eggshells vs. without eggshells. We also experimentally examined the effects of egg aggregation by incubating eggs singly, in groups of four, and in groups of nine. We recorded egg surface temperature, water uptake, and hatchling morphology. Females were more likely to nest in pots with used soil and with eggshells than in pots with fresh soil or without eggshells. We observed no effects of egg aggregation on egg survival, egg temperature, or most measures of hatchling morphology. However, singly-incubated eggs absorbed more water than eggs incubated in the four and nine egg aggregations and this resulted in offspring with greater body condition (i.e. heavier for their length) at hatching. The behavioural experiment demonstrates that females actively choose nest sites that have been used previously (as expected under the adaptive hypothesis), but the egg-aggregation experiment shows no benefits to offspring based on the variables measured. Thus, results of the behaviour study support the adaptive hypothesis; however, results from our egg-incubation study do not. Likely, the adaptive and constraint hypotheses are not mutually exclusive, and a diversity of factors influence the evolution of communal nesting behaviour.     

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.     

The exclusion of avian predators from aggregations of nesting lapwings (Vanellus vanellus)

Territorial lapwings in Aberdeenshire, Scotland, largely prevented carrion crows, (Corvus corone), their main egg predators, from approaching close to nest-sites and from entering the area within an aggregation of 11 nests. Predation rates of artifical nests were significantly lower when placed within 10 m of active lapwing nests than when>200 m from nest-sites, indicating that the presence of lapwings afforded protection from crow predation. Further artificial nest experiments showed that the amount of protection declined linearly with increasing distance from the nest-site, and some degree of protection occurred to at least 30–50 m from it. The size of this protected zone was apparently related to the number of lapwing nests present, and the survival time of eggs in artificial nests reflected the lapwings' nesting density. The defended zones around each lapwing nest overlapped appreciably in dense or large nesting aggregations, leading to the observed protection by generally excluding crows, from the nesting area.     

Social Behaviour of the Carpenter Bee Xylocopa pubescens (Spinola)

The development of about 20 relatively small nests of Xylocopa pubescens was studied. After the first offspring had become adult, these nests reached a social stage in which there was only one egg-layer per nest. Freshly emerged (teneral) adults eat a lot of food, collected by a forager, before they fly out of the nest. This food appears to be of major importance to these bees in that it makes them fully agile. It seems therefore, that part of the food needed for the development of a young bee is not given at the larval stage, via the beebread, but at the teneral adult stage. As a result of this, a necessary overlap of generations is accomplished. Besides, less pollen has to be collected for the provision of a brood cell, so more cells can be constructed within a short period of time. There is a high degree of nest competition. Many nests were taken over by conspecific individuals (or by females of X. sulcatipes) that were searching for nesting sites within the study area. However, although more than 50% of the solitary nests were taken over sooner or later, strangers hardly ever intruded into social nests, with more than one adult. This illustrates how important it is for a reproducing female to tolerate the presence of nestmates which guard the nest in her absence. Although in most cases the foundress of a nest proceeded to produce new brood in the presence of her offspring, it happened that a daughter took over reproduction from her mother. In two of these cases, it could be observed that a mother started a new nest elsewhere after having been thrown out by her own daughter. At least in these cases, nest competition between mother and daughter started long before the mother had reached the end of her reproductive capacities. Since nesting possibilities are scarce, it seems a logical strategy to stay in the maternal nest and wait for a chance to become the egg-layer if the mother dies or if she loses her dominance. The occurrence of several social interactions among nestmates is discussed: — trophallaxis was observed often, not only under ‘forced’ conditions, but also as a form of ‘voluntary’ feeding; — nestmates were observed to groom each other.     

Group nesting and individual variation in behavior and physiology in the orchid bee <Emphasis Type="Italic">Euglossa nigropilosa</Emphasis> Moure (Hymenoptera,Apidae)

The Euglossini are a key group for studying the traits that promote or hinder highly social behavior in bees because it is the only tribe in the Apine clade without large colonies or females with distinct life histories, e.g. queens and workers. There have been few studies on behavior of orchid bee females in nests because these nests are not found easily. Taking advantage of the relatively high abundance of Eg. nigropilosa nests at Reserva Natural La Planada, Colombia, we examined social behavior of Eg. nigropilosa individuals in five nests (3 original and 2 reused) for nine months. We report this species to have the largest colonies known for Euglossa, with nests reaching up to 22 individuals, and all nests containing more than one female bee from the same generation. These nests presented many traits that correspond to communal insect colonies. No generational overlap and no cooperative brood care were detected. We examined natural enemies and resource limitation as important factors for group nesting. We examined parasitoid attacks to cells in a nest with females and one without females. We also searched for nesting locations and examined nest re-use as indicators of nest site limitation. Lastly, we examined behavioral and physiological differences among females in the same nest. Such differences could be the bases for evolution of alternative life histories among group living females. We examined extent of ovary development and oviposition rates in similarly aged females in the same nest. We found large variation in reproductive effort of young females. We also examined differences in resin foraging and cell usurpation behaviors. Behavioral specialization was observed, with some individuals bringing only resin to the nest. Inside the nests, bees had territories in which they constructed and defended cells. This territoriality may be a defense against usurpation of provisioned cells by nest mates. Received 10 December 2007; revised 2 May 2008; accepted 7 May 2008.     

Predator–prey interactions between the South Polar skua Catharacta maccormicki and Antarctic tern Sterna vittata

Antarctic terns have to co‐exist in a limited space with their major nest predator, the skuas. We conducted artificial nest experiments to evaluate the roles of parental activity, nest location and nest and egg crypsis in this simple predator–prey system. Predation on artificial (inactive) nests was higher in traditional nesting sites than in sites previously not occupied by terns, which suggests that skuas memorized past tern breeding sites. Predation on artificial nests in inactive colonies was higher than in active (defended) colonies. Parental defense reduced predation in colonies to the level observed in artificial nests placed away from colonies. This suggests that communal defense can balance the costs of attracting predators to active colonies. Within colonies, predation was marginally higher on experimental eggs put in real nests than on bare ground. Although it seems that the presence of a nest is costly in terms of increased predation, reductions in nest size might be constrained by the need for protective nest structures and/or balanced by opposing selection on nest size. Predation did not differ markedly between artificial (quail) and real tern eggs. A simultaneous prey choice experiment showed that the observed predation rates reflected egg/nest detectability, rather than discrimination of egg types. In summary, nesting terns probably cannot avoid being detected, and they cannot defend their nest by attending them. Yet, by temporarily leaving the nest, they can defend it through communal predator mobbing, and at the same time, they can benefit from crypsis of unattended nest and eggs.