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.     

Nesting success of wood‐cavity‐nesting bees declines with increasing time since wildfire

Bees require distinct foraging and nesting resources to occur in close proximity. However, spatial and temporal patterns in the availability and quantity of these resources can be affected by disturbances like wildfire. The potential for spatial or temporal separation of foraging and nesting resources is of particular concern for solitary wood‐cavity‐nesting bees as they are central‐place, short‐distance foragers once they have established their nest. Often the importance of nesting resources for bees have been tested by sampling foraging bees as a proxy, and nesting bees have rarely been studied in a community context, particularly postdisturbance. We tested how wood‐cavity‐nesting bee species richness, nesting success, and nesting and floral resources varied across gradients of wildfire severity and time‐since‐burn. We sampled nesting bees via nesting boxes within four wildfires in southwest Montana, USA, using a space‐for‐time substitution chronosequence approach spanning 3–25 years postburn and including an unburned control. We found that bee nesting success and species richness declined with increasing time postburn, with a complete lack of successful bee nesting in unburned areas. Nesting and floral resources were highly variable across both burn severity and time‐since‐burn, yet generally did not have strong effects on nesting success. Our results together suggest that burned areas may provide important habitat for wood‐cavity‐nesting bees in this system. Given ongoing fire regime shifts as well as other threats facing wild bee communities, this work helps provide essential information necessary for the management and conservation of wood‐cavity‐nesting bees.     

Review of bees as guests in termite nests,with a new record of the communal bee, <Emphasis Type="Italic">Gaesochira obscura</Emphasis> (Smith, 1879) (Hymenoptera,Apidae), in nests of <Emphasis Type="Italic">Anoplotermes banksi</Emphasis> Emerson, 1925 (Isoptera,Termitidae, Apicotermitinae)

Associations between bees and termites are documented infrequently, but records are available for bee species ranging in behavior from solitary to highly eusocial. The subtribe Meliponina (stingless bees) is the most common bee group reported in termite nests, and, for some species, the occupancy of termite nests may be obligatory. The records of solitary bees nesting within termite nests include species of the tribes Emphorini, Centridini, Megachilini, and Paracolletini. Most of these bees can probably nest in other substrates, and their relationships with termite nests are presumably opportunistic. This study provides a review of published records of bees as guests in termite nests, and also describes the aggregation of nests of Gaesochira obscura within one nest of Anoplotermes banksi in Brazilian Amazonia. One termite nest contained at least ten nest entrances of G. obscura, with burrows 4–6 mm in diameter and about 10 cm long. Each burrow ended in brood cells in different stages of food provisioning and larval development. As commonly reported for other associations of this nature, there was no connection between the tunnels of bees and those of termites. This record adds important data on the biology of A. banksi. Because this is a single record, it is impossible to classify G. obscura either as a termitophile or termitariophile; this species may be opportunistic in relation to nesting substrate.     

Fitness consequences of ecological constraints and implications for the evolution of sociality in an incipiently social bee

Ecological constraints such as resource limitation, unfavourable weather conditions, and parasite pressure have long been considered some of the most important selective pressures for the evolution of sociality. In the present study, we assess the fitness consequences of these three ecological factors on reproductive success of solitary nests and social colonies in the socially polymorphic small carpenter bee, Ceratina australensis, based on 982 nests collected over four reproductive periods. Nest site limitation was predicted to decrease opportunities for independent nest initiation and increase the frequency of social nesting. Nest sites were not limiting in this species and the frequency of social nesting was consistent across the four brood‐rearing periods studied. Unfavourable weather was predicted to lower the frequency of female dispersal from their natal nests and to limit the brood‐rearing season; this would increase the frequency and fitness of social colonies. Daily temperature and precipitation accumulation varied between seasons but were not correlated with reproductive success in this bee. Increased parasite pressure is predicted to increase the frequency and fitness of social colonies because solitary bees must leave the nest unattended during foraging bouts and are less able to defend the nest against parasites. Severe parasitism by a chalcid wasp (Eurytoma sp.) resulted in low reproductive success and total nest failure in solitary nests. Social colonies had higher reproductive success and were never extirpated by parasites. The high frequency of solitary nests suggests that this is the optimal strategy. However, social colonies have a selective advantage over solitary nesting females during periods of extreme parasite pressure, and we suggest that social nesting represents a form of bet‐hedging against unpredictable fluctuations in parasite number. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 57–67.     

Bee Tracker—an open‐source machine learning‐based video analysis software for the assessment of nesting and foraging performance of cavity‐nesting solitary bees

The foraging and nesting performance of bees can provide important information on bee health and is of interest for risk and impact assessment of environmental stressors. While radiofrequency identification (RFID) technology is an efficient tool increasingly used for the collection of behavioral data in social bee species such as honeybees, behavioral studies on solitary bees still largely depend on direct observations, which is very time‐consuming. Here, we present a novel automated methodological approach of individually and simultaneously tracking and analyzing foraging and nesting behavior of numerous cavity‐nesting solitary bees. The approach consists of monitoring nesting units by video recording and automated analysis of videos by machine learning‐based software. This Bee Tracker software consists of four trained deep learning networks to detect bees that enter or leave their nest and to recognize individual IDs on the bees’ thorax and the IDs of their nests according to their positions in the nesting unit. The software is able to identify each nest of each individual nesting bee, which permits to measure individual‐based measures of reproductive success. Moreover, the software quantifies the number of cavities a female enters until it finds its nest as a proxy of nest recognition, and it provides information on the number and duration of foraging trips. By training the software on 8 videos recording 24 nesting females per video, the software achieved a precision of 96% correct measurements of these parameters. The software could be adapted to various experimental setups by training it according to a set of videos. The presented method allows to efficiently collect large amounts of data on cavity‐nesting solitary bee species and represents a promising new tool for the monitoring and assessment of behavior and reproductive success under laboratory, semi‐field, and field conditions.     

Nesting activity of cavity-nesting bees and wasps is lower in small-scale apple orchards compared to nearby semi-natural habitats

1. Commercially reared cavity-nesting bees have been studied mainly in large, intensively managed orchards. However, knowledge on wild cavity-nesting bee and wasp communities and their potential limitations in smaller orchards remain insufficient.
2. We compared the colonization rate of trapnests, nesting success, parasitism and response to flower resources of cavity-nesting bees and wasps between apple orchards and nearby semi-natural habitats (SNHs).
3. Trapnests were placed in orchards and neighbouring SNHs. Colonization dynamics were studied and herbaceous flower resources were estimated. Furthermore, nest and brood cell quantity, number of alive offspring and nest parasitism rate were assessed.
4. We found a higher colonization rate in the SNHs than in the orchards. Both bees and wasps made more nests, completed more brood cells and had a higher number of alive offspring in the SNHs. The number of bee nests in the orchards showed a positive correlation with the species richness of the flowering plants. The nest parasitism of wasps was higher in the SNHs.
5. Apple orchards in the studied small-scale system were generally less colonized by cavity-nesting hymenopterans than nearby SNHs that can be important reservoirs of these ecosystem service provider hymenopterans. Our results highlight the importance of diverse flowering herbaceous vegetation in the understory that increased the number of bee nests in orchards and that could have a positive effect on the nesting activity of the bee species active in summer. Therefore, management practices that support flowering plant species in the understory vegetation are highly recommended in such orchards.

     

Biology of the mason bee Osmia latreillei (Hymenoptera: Megachilidae) under artificial nesting conditions in Egypt

The mason bee Osmia (Helicosmia) latreillei Spinola is one of the dominant species of bees throughout Egypt, and can be found commonly in the Suez Canal Region. The species visits numerous plant species of the family Asteraceae and is considered the most important pollinator of certain crops such as sunflower. This species and some other solitary cavity nesting bees are threatened by fragmentation of their nesting habitats. Several attempts were made to establish and propagate O. latreillei, and success was achieved for re-nesting it in various artificial materials such as wood and polystyrene with rolled paper straws. These artificial nests were established at the conservation area of the Bee Research Centre, Suez Canal University, in order to study the nesting behavior and biology of O. latreillei under artificial conditions and to document their natural enemies. Biologically, there was no significance difference between the life cycle of O. latreillei under natural and artificial nesting conditions. This solitary bee was univoltine, individuals active only during Spring, and attacked by diverse enemies, with Stelis (Stelis) murina Pérez and chrysidid wasps considered the two most major pests of this wild bee species.     

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.     

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.     

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.     

Colony disturbance and solitary nest initiation by workers in the obligately eusocial sweat bee, Halictus ligatus

Sweat bees are one of the most socially polymorphic lineages on the planet. In obligately eusocial species, newly enclosed females may become either queens or workers, depending on the environmental and social circumstances of the nest into which they emerge. In socially polymorphic species, females also have the option of nesting solitarily, founding a nest and raising future reproductives alone, without the help of other adult females. Halictus ligatus is a widespread Nearctic, ground-nesting sweat bee. It has been particularly well studied in Ontario, where detailed studies have described it as obligately eusocial. Here we report evidence that the flexibility of female H. ligatus actually extends to expressing behaviour more typical of socially polymorphic species, those in which some individuals reproduce solitarily. In a population in southern Ontario, black wasps (Astata sp.) emerged from the soil beneath the nesting aggregation and proceeded to excavate their own nesting tunnels, dislocating many H. ligatus nest entrances. Young workers whose natal nests were destroyed by the wasp activity constructed new nests, so under very specific circumstances, it is possible for potential altruists to nest solitarily.     

Trap-nest occupation by solitary wasps and bees (Hymenoptera: Aculeata) in a forest urban remanent

Temporal variation of solitary wasps and bees, nesting frequency, mortality, and parasitism were recorded from a remanent forest in Belo Horizonte, MG, Brazil. Wasps and bees were collected in trap-nests placed in areas with 25, 100, and 400 m2, from February to November 2004. The 137 trap-nests collected contained 11 species of wasps and bees. Wasps occupied most nests (75%). Occupation peaks occurred in March (25%) and September (26%); in June, the lowest occupation (2%) was observed. Except for Trypoxylon (Trypargilum) lactitarse Saussure, no significant correlation was found between number of occupied nests, and temperature and rainfall means. In the nests, 48% of the immature specimens died; 13% of the nests were parasitized. Total death and parasitism rates of wasps and bees differed significantly.     

Influence of the Orientation of Nest Entrance, Shading, and Substrate on Sampling Trap-Nesting Bees and Wasps

The use of trap nests supplies important biological information concerning the biology of bees and wasps. However, the knowledge about the factors that influence the management of these groups is still scarce. Our study aimed to analyze the effects of the orientation of nest entrance, shading, and block material (wood or vermiculite) on the frequency, richness, and composition of trap-nesting species at two agricultural areas in the Atlantic Forest region of the state of Paraiba. Both sites showed similar species richness and composition. However, a higher number of nests was found at the Alhandra site, mainly of the species Trypoxylon aurifrons Shuckard and Centris analis (F.). Regarding the orientation, the number of nest entrances oriented to prevailing wind direction was significantly smaller. Most species nested more frequently in shaded areas than in sunny areas and the wood blocks were more attractive than the vermiculite blocks, especially for bees.     

Nest preference and ecology of cavity-nesting bees (Hymenoptera: Apoidea) in Punjab,Pakistan

Among bees, 85 % are solitary species, most of them are ground-nester and some are cavity-nesting and construct their nests in pre-existing cavities. This work was conducted to evaluate the substrate preference and nest architecture (acceptance, occupation percentage, seasonality and parasitism) of cavity-nesting bees in different substrates. Trap nests offered comprised five different materials (drilled cavities in wood, bamboo, cardboard tubes, plastic soda straws, and mud blocks). These were installed in four districts for two years. The nesting cavities of five different diameters (6, 8, 10, 12 and 14 mm) were provided in each nesting material with an average length of 180 ± 9.92 mm. In all the materials, 5400 nesting cavities were offered, out of which 628 were colonized by bees from two families (Megachilidae and Apidae) and six species (Megachile cephalotes, M. lanata, M. bicolor, Xylocopa basalis, X. fenestrata and Ceratina smaragdula), including one parasitic bee (Euaspis carbonaria). The bee species differed significantly in occupying five nesting materials. The most preferred diameters were 8 mm and 10 mm, with 52.20% and 29.45% of colonization, respectively. Nesting was done throughout the year except in winter. This study will serve as a baseline for future studies and conservation programs of cavity-nesting bees in Pakistan.     

木蜂的巢内行为和交配策略

木蜂Xylocopa作为蜜蜂科的一个重要类群, 用来研究昆虫社会进化早期阶段具有重要意义。本文综述了近年来木蜂的营巢习性、 社会性行为和交配策略的研究进展。木蜂巢的建筑与巢内巢室的分布、 巢室的大小、 贮备蜂粮的效率和被寄生的敏感性等均有一定的关系。在筑巢地点, 随着木蜂种群数量的增加, 同种雌蜂之间的资源存在着竞争, 木蜂对巢室的守卫加强, 更多地表现为对同种雌蜂的守卫以及防御其他的天敌入侵。其社会多态性由独栖性向社会性演化, 主要表现为独栖性、 亚社会性、 半社会性、 共生性和准社会性等方式, 甚至同一种群内会出现不同的社会性行为。在交配策略上, 木蜂的雄蜂在外部形态特征上具有适应交配和寻找雌蜂的进化特征, 并且具有复杂的领地行为。这些研究理论对于我们深刻理解木蜂的行为生物学具有重要意义, 同时也有助于进行更深层次的社会性蜂类演化进程的探索。     

‘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.     

Cavity-nesting bees and wasps (Hymenoptera: Aculeata) in a semi-deciduous Atlantic forest fragment immersed in a matrix of agricultural land

Cavity-nesting bees and wasps (Hymenoptera: Aculeata) have been showed to be suitable models to investigate the effects of forest fragmentation and human land use. Those studies are particularly pertinent when considering fragmented ecosystems such as the Atlantic semi-deciduous forest in Brazil. We investigated the changes in composition, nest abundance, and mortality of cavity-nesting bees and wasps over edge-center forest fragment immersed in a matrix of agricultural lands in southeastern Brazil. Trap-nests (bamboo canes and cardboard tubes) were set in nine sampling stations in three different zones in the study site: three sampling stations at the forest edge; three at 250 m away from the edge and another three at 500 m away from the edge. Nests were monitored monthly for 2 years (from June 2011 to May 2013). A total of 942 nests (706 built by 16 bee species; 236 from 18 wasp species) were collected in the fragment. A significant change over gradient edge-center was observed on the parameters analyzed. The Non-metric Multidimensional Scale analysis showed that 72% of the species (host and natural enemies) that present more than ten records were associated with the forest edge. Furthermore, the highest values of the abundance of nests, parasitism, and mortality were recorded in the edge. In spite of being surrounded by farmlands, the fragment hosts a great diversity of trap-nesting wasp and bee species. Our findings demonstrate that forest edges are important habitats to maintain communities of cavity-nesting bees and wasps.     

Spillover of trap-nesting bees and wasps in an urban–rural interface

A mismatch of resource availability in certain periods can lead to spillover of insects between habitats, resulting in temporal differences in insect diversity. Urban gardens are important anthropogenic habitats but it is unknown whether, when and why spillover of beneficial insects occurs between gardens and agricultural habitats. We used trap nests for Hymenoptera to monthly monitor bee and wasp abundance and species richness in 12 gardens and 12 rapeseed fields. Half of the gardens and rapeseed fields were located in the urban–rural interface and bordered each other (a garden paired with a rapeseed field) and the other half were isolated in the rural landscape (isolated rapeseed fields) and in the urban city centre (isolated gardens). In general, gardens in the urban–rural interface comprised the highest richness of bees and wasps. The abundance of bees but not of wasps was highest in paired habitats and peaked at full rapeseed blooming, indicating that mass-flowering rapeseed offers foraging resources for bees nesting in adjacent gardens. Thus, bees nest and increase their populations in both areas, benefiting from the mass-flowering resource in the agricultural habitat as well as the nesting resources from gardens, suggesting spillover of bees but not of wasps between paired gardens and rapeseed fields. Our study highlights the value of gardens in the urban–rural interface for the biodiversity of functionally important insects. Implementing urban gardening and small-scale agriculture in cities and suburban habitats can promote local pollinator populations and benefit adjacent croplands.     

Why Wasp Foundresses Change Nests: Relatedness,Dominance, and Nest Quality

The costs and benefits of different social options are best understood when individuals can be followed as they make different choices, something that can be difficult in social insects. In this detailed study, we follow overwintered females of the social wasp Polistes carolina through different nesting strategies in a stratified habitat where nest site quality varies with proximity to a foraging area, and genetic relatedness among females is known. Females may initiate nests, join nests temporarily or permanently, or abandon nests. Females can become helpers or egglayers, effectively workers or queens. What they actually do can be predicted by a combination of ecological and relatedness factors. Advantages through increased lifetime success of individuals and nests drives foundresses of the social wasp Polistes from solitary to social nest founding. We studied reproductive options of spring foundresses of P. carolina by monitoring individually-marked wasps and assessing reproductive success of each foundress by using DNA microsatellites. We examined what behavioral decisions foundresses make after relaxing a strong ecological constraint, shortage of nesting sites. We also look at the reproductive consequences of different behaviors. As in other Polistes, the most successful strategy for a foundress was to initiate a nest as early as possible and then accept others as subordinates. A common feature for many P. carolina foundresses was, however, that they reassessed their reproductive options by actively monitoring other nests at the field site and sometimes moving permanently to new nests should that offer better (inclusive) fitness prospects compared to their original nests. A clear motivation for moving to new nests was high genetic relatedness; by the end of the foundress period all females were on nests with full sisters.     

An experimental analysis of joint nesting in the salamander Hemidaetylium scutatum (Caudata: Plethodontidae): the effects of population density

Three nesting behaviour patterns are documented in the plethodontid salamander Hemidactylium scutatum. A female may lay eggs (1) in a solitary nest and brood them, (2) in a joint nest and brood them as well as eggs of other females, or (3) in a joint nest that is brooded by another female. The hypothesis that population density was positively associated with joint nesting was tested by following two populations for 5 years and by experimentally manipulating the population density of nesting females in artificial habitats for the latter 2 years. The proportion of joint nests did not vary with density, although joint nests tended to contain eggs of more females at the high population density. Joint nests were usually brooded by one female; thus, most females that laid eggs in joint nests did not brood them at high density. The reproductive success, as measured by survival of embryos, of solitary and joint nesters was equivalent. Joint nests were deserted less often, however, which decreased the probability of catastrophic mortality. The number of days of brooding was significantly positively correlated with loss of body mass of females, suggesting a cost to brooding behaviour. Joint nesting with solitary brooding is not explained by aggressive usurpation of nests or by brood parasitism.