Nest‐to‐nest dispersal of Chaetodactylus krombeini (Acari,Chaetodactylidae) associated with Osmia cornifrons (Hym., Megachilidae)

A cleptoparasitic mite, the Krombein’s hairy‐footed mite, Chaetodactylus krombeini Baker (Acari, Chaetodactylidae) became a key pest that affects the maintenance and propagation of Osmia spp. (Hym., Megachilidae), thus disrupting orchard pollination in the United States. Although hypopi, the dispersal stages of C. krombeini, are known to disperse from nest to nest by hitchhiking on Osmia cornifrons adults, we observed that they might disperse in other ways too in commercial orchards. This study was conducted to elucidate the nest‐to‐nest dispersal mechanisms of C. krombeini hypopi. We tested three potential dispersal mechanisms of C. krombeini other than phoresy by O. cornifrons: (1) dispersal by walking from nest to entrances of nearby nests, (2) dispersal by walking from nest to nest through emergence holes made by parasitic wasps on nests, and (3) dispersal by being unloaded and uptaken to and from flowers by O. cornifrons. Results of this study showed that C. krombeini hypopi could disperse from a nest to nearby nests by walking through nest entrances and holes made by parasitic wasps of O. cornifrons. Although 0.06% of C. krombeini hypopi on blueberry flowers were picked up by O. cornifrons, they were not able to be unloaded to flowers from O. cornifrons and no hypopi could inhabit or survive on blueberry flowers. This indicated no or very low chance of C. krombeini hypopi dispersal via blueberry flowers. Based on our findings of C. krombeini dispersal ecology, development of C. krombeini control strategies are discussed in this article.     

Modeling spring emergence of Osmia cornifrons Radoszkowski (Hymenoptera: Megachilidae) females in Korea

The Japanese hornfaced bee, Osmia cornifrons Radoszkowski (Hymenoptera: Megachilidae), is an important pollinator of apple. It overwinters as diapausing adult in cocoon and the synchronization of the adult emergence in spring with apple blossom is critical for successful pollination, specifically in the environment with variable climate. To this end, prediction model of adult emergence would help manage O. cornifrons populations for planed pollination. In this study, by using temperature-dependent emergence data of O. cornifrons, emergence rate were mathematically described using linear and Lactin's nonlinear regression methods. The lower emergence thresholds estimated from linear regression for female was 7.98 °C and the thermal constant (DD, degree days) was 112.43DD. Pattern of emergence completion was described by using Weibull function and prediction of spring emergence was modeled based on functions of emergence rate and distribution. The model predicted the emergence of O. cornifrons between 100–130 Julian date and the observation fell between 108–129 Julian date. The prediction ranges of O. cornifrons emergence were wider than that of observation. This model could help predict spring emergence of O. cornifrons in the field and optimize planed pollination program for pollination service in agricultural systems in the climate change era.     

Within-nest spawning-site preferences of female bluefin damselfish: the effect of early-stage eggs

In some fish species with exclusive paternal care females prefer to deposit their eggs in nests containing early-stage eggs and avoid those containing only late-stage eggs. Such female preference may also be shown when it comes to deciding where to lay within a given nest; that is, females may prefer to lay contiguously to younger rather than to older eggs. To understand if females benefit from such preferences by obtaining higher offspring survival, we conducted field studies with the bluefin damselfish, Abudefduf luridus. We investigated (1) female between-nest and within-nest spawning-site preferences relative to brood characters and (2) egg-hatching success related to male filial cannibalism. We found evidence that males entirely ate some of their clutches, especially the eggs received near the end of the breeding season. These were also more susceptible to being eaten when laid in nests containing only eggs in a late stage of development. Theoretically, these patterns of clutch loss should result in adaptive female preference for nests containing early-stage eggs, but we found no evidence for such a preference in female A. luridus. However, once females had chosen a nest they strongly preferred to lay contiguously to early-stage eggs than to late-stage eggs. Moreover, females benefited from this decision by obtaining higher survival rates for their clutches. We conclude that within-nest spawning-site preference of the female for early-stage eggs may have evolved as an adaptive response to a dilution effect on the risk of egg loss.     

The nesting biology of the bee, Osmia dimidiata Morawitz, 1870 (Hymenoptera, Megachilidae) in the Crimea

The structure and composition of Osmia dimidiata nests were studied based on the material of 133 nests obtained from trap-nests in five localities of the Crimea. The nests contained from 1 to 9 cells (on average, 2.2), separated with partitions made of masticated leaves. Most of the cells had a massive hemispherical bottom and a thin flat cover. The final plug had a structure similar to the cell bottom. The cells occurred singly in the nest cavity or were arranged in rows of 2 or 3 cells separated by galleries. The diameter of the occupied cavities was positively correlated with the fraction of the females in the progeny and with the mass of the cocoons, but negatively correlated with the cell length. The cells with females were 1.5 times as long as those with males. The cocoons with females were twice as heavy as those with males. The peculiar characters of the nest structure of O. dimidiata may mean that the typical nesting substrates of this species are abandoned cells of other bee and wasp species. The characteristic mechanism of estimating the mass of the pollen loaf stored in the cells and determining the brood sex ratio has evolved in the O. dimidiata due to its tenantry. This mechanism is based on two constants: the sex-linked cell length and the fixed distance between the pollen loaf and the cell cover. When choosing the nesting cavities, the female prefers broader and longer tubes for cells with females and narrower and shorter ones for cells with males. The overall sex ratio of the progeny was male-biased (1 ♀: 2.5 ♂) and depended on the food supply abundance. The adults of O. dimidiata fly in the Crimea from the last decade of May until the end of July. Hibernation occurs at the adult stage in the cocoons. Females collect pollen from flowers of Asteraceae or, rarely, Fabaceae. The nests may be destroyed by Melittobia acasta, Monodontomerus aereus, M. obscurus, Sapyga quinquepunctata, Stelis phaeoptera, and Trichodes apiarius; 21% of the progeny died from these enemies, and 22%, due to other factors. The previously proposed synonymy of Pseudosmia taurica Radoszkowski, 1874 (nec Osmia taurica Radoszkowski, 1887) and O. dimidiata is rejected.     

Life cycle of Anthracophora rusticola (Coleoptera: Cetoniidae) from the commensal nests of Chinese Sparrowhawks (Accipiter soloensis)

Commensal breeding habits of the Anthracophora rusticola (Coleoptera: Cetoniidae) in stick nests of carnivorous, piscivorous, and omnivorous birds were recently reported, but the detailed life history of the scarab beetle has not been thoroughly elucidated. To understand the breeding cycle of the declining A. rusticola related to that of nesting Chinese sparrowhawks (Accipiter soloensis) in Korea, we examined six sparrowhawk nests from 2005 to 2008 and reared larvae and cocoons in a cage in 2008. Each sparrowhawk nest hosted 11.3 ± 3.0 larvae and cocoons, implying that sparrowhawk nests might be the most important nesting resource of the beetles in Korea. The beetles may lay their eggs between late May and early July after nest building by sparrowhawks. Larvae were observed from July to August in the sparrowhawk nests, and adult beetles emerged in September after a mean pupal period of 25.5 days. Our results on the coinciding breeding of two commensal taxa over three months from May to September may provide key information regarding the life history of the commensal scarabid beetle and potential causes of the concurrent decline of both taxa in Korea.     

Female within-nest spawning-site preference in a paternal brooding blenny and its effect on the female mate choice

Female spawning-site preference within a nest and its effect on the female mate choice in a paternal brooding blenny Rhabdoblennius ellipes , were examined in rocky intertidal pools using artificial nests. The number of eggs deposited at the nest entrance site was lower than the number deposited at the center and at deep sites. Moreover, the within-nest difference in the egg developmental stage indicated that eggs at the nest entrance site were deposited after those at the center and deep sites. These results indicated that females prefer to spawn eggs in the center and deep sites rather than at the entrance site. Owing to the higher egg mortality rate at the entrance site, females may avoid spawning at such sites. An analysis of the nests in the study area suggested that the within-nest site preference affects female mate choice that is females avoided nests where only the entrance site was available for spawning and instead spawned in nests where the center and deep sites were available.     

Foraging and nesting behavior of Osmia lignaria (Hymenoptera: Megachilidae) in the presence of fungicides: cage studies

During orchard pollination studies in California, we observed dramatic changes in nesting and foraging behavior of Osmia lignaria Say (Hymenoptera: Megachilidae) after sprays with tank mixtures containing fungicides. A characteristic pattern of postspray events observed includes erratic behavior and interrupted foraging and nesting activity for several days. In an effort to determine whether fungicidal sprays were disruptive to bee foraging and thus to pollination, we exposed O. lignaria females nesting in field cages planted with lacy scorpionweed, Phacelia tanacetifolia Benth (Hydrophyllaceae), to selected spray mixtures normally encountered in California orchard production systems: iprodione (Rovral), propiconazole (Orbit), benomyl (Benlate), and captan (Captan 50 WP); the surfactant Dyne-Amic, alone and mixed with Rovral; and the tank mixture IDB (Rovral + Dyne-Amic + the foliar fertilizer Bayfolan Plus). An additional cage sprayed with an equal volume of water acted as control, and a cage sprayed with the insecticide dimethoate as a toxic standard. For each female O. lignaria, we recorded time spent inside the nest depositing pollen-nectar loads, foraging time, cell production rate, and survival. All females in the dimethoate treatment died postspray + 1 d. Before death, some of these females behaved similarly to our previous orchard observations. A high proportion of females in the IDB cage were inactive for a few hours before resuming normal foraging and nesting activity. No lethal or behavioral effects were found for any of the other compounds or mixtures tested. Our results indicate that the fungicide applications that we tested are compatible with the use of O. lignaria as an orchard pollinator.     

Strongly biased sex ratio in cuckoo wasp Chrysura hirsuta (Hymenoptera: Chrysididae), a parasitoid of the mason bee Osmia orientalis

We examined the rate of parasitism and sex ratio of the cuckoo wasp Chrysura hirsuta (Gerstaecker) (Hymenoptera: Chrysididae) that emerged from nests of the mason bee Osmia orientalis Benoist (Hymenoptera: Megachilidae) in Nara, Japan. Nests of O. orientalis were found in empty shells of two snail species, Satsuma japonica (Pfeiffer) and Euhadra amaliae (Kobelt). The percentage of parasitism by cuckoo wasps per all collected cocoons tended to be high (20–50%) even though interannual variation and the average number of cocoons per nest did not differ across snail shell species within each year. Our results from three years of observation, combined with previous reports, showed that the adult sex ratio of C. hirsuta was strongly female‐biased, which suggests that the species reproduces by thelytokous parthenogenesis.     

The nest structure in four wasp species of the genus Euodynerus Dalla Torre (Hymenoptera, Vespidae: Eumeninae)

A comparative analysis of the main parameters of the nest structure and composition in Euodynerus quadrifasciatus, Eu. dantici, Eu. disconotatus, and Eu. velutinus in the Crimea is given. Nests of all the studied species were obtained from trap-nests made of reed stems; the nests of Eu. dantici and Eu. disconotatus were additionally obtained from the abandoned cells of the nests of Sceliphron destillatorium. The nest of Eu. quadrifasciatus consists of a consecutive row of cells, each cell having its own bottom and lid with empty space between them; the nests of three other species consist of an uninterrupted row of cells without spaces, so that the cell bottoms act as partitions. The data on the diameter and length of nest cavities occupied by the studied species, the number of cells in nests, the sex ratio and the length of the cells with prospective females and males are reported. The correlations between the length of the cells and the diameter of the occupied cavities and between the length of the rear empty nest space and vestibule and the length of the occupied nest cavity are analyzed. The structure of cocoons and final nest plugs are described. The difference in the evolution of nest building instincts in members of the subgenera Pareuodynerus and Euodynerus s. str. and the role of adaptation to nest tenantry in the evolution of nest building instincts of Eu. dantici and Eu. disconotatus are discussed.     

Redescription of Natalaspis leptocarpi n.comb. (Homoptera: Diaspididae)

Abstract

Cyllobelus rufopictus? a salticid species from equatorial Africa, has a large and complex repertoire of displays. These spiders live on and near the ground in habitats of short grass, and build opaque silk nests on dead leaves. Each male uses one of three different mating tactics depending on the female’s maturity and location (communicatory versatility). Type 1 courtship, involving special movements and postures of the legs, palps, and body, occurs if the female is an adult away from the nest; apparently this type of courtship is vision-dependent. If the male encounters an adult female inside her nest, he uses Type 2 courtship, which consists of movements by the male that cause the silk to vibrate. If the female is a subadult inside her nest, the male initially uses Type 2 courtship, then builds a second chamber on the female’s nest, and cohabits until she moults and matures. Other displays occur during male-male and female-female interactions. Biting and pinching are distinctive Type 2 courtship displays that have not been described previously for salticids, and juddering is the most distinctive Type 1 display.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Big is better: the importance of size for reproductive success in male Pomatoschistus minutus (Pallas) (Pisces, Gobiidae)

The sand goby, Pomatoschistus minutus , was used in aquarium experiments to study the importance of body size for the ability of males to gain access to nest sites and mates. When several male and female P. minutus were allowed to reproduce together, on average, half of the males built nests, and half of those males obtained eggs in their nests. Males with nests were significantly larger than males without nests, and nests with eggs belonged to males larger than the males with empty nests. In another experiment, when two males were competing for one nest, the largest male occupied the nest when both fish were put into the aquarium simultaneously. However, when the smaller male had been allowed to establish a nest before the larger male was introduced, the small male could usually retain the nest. Males with a body length < 50 mm did not build nests at all in the early part of the breeding season. In female choice experiments, no preference for larger males was found. Thus, male-male competition for nest sites and behavioural differences between different sized males seem to be the main factors influencing the non-random mating success in male P. minutus.     

Nesting patterns of European hedgehogs (Erinaceus europaeus) under northern conditions

Little information exists on the nesting of hedgehogs (Erinaceus europaeus) in the harsh northern environment, although this aspect has specific importance for the species’ survival in this region. We studied the nest types and nesting site selection of radio tagged hedgehogs in urban areas in Eastern Finland. Altogether, four different nest types were discovered and one of them (the pre-hibernation nest) is described here for the first time. Hedgehogs preferred residential areas for nesting during the mating and post-mating seasons while in the pre-hibernation season and hibernation period nesting was more common in forest areas. Nest constructions were closely related to materials found in the nest vicinity. During the active period, males used on average 24 nests and changed them 30 times, whereas females used 17 nests and changed them 26 times. The number of female nests and frequency of nest change increased from post-mating to pre-hibernation seasons, while in males both decreased. In general, female and male nests were situated within their established long-term home range, but in the pre-hibernation season over 30 % of male nests were situated outside of it. In post-mating and pre-hibernation seasons, females avoided other individuals of the same sex; this was demonstrated by the lack of nests in other female core areas. Hedgehogs hibernate for around 223 days, which is the longest reported hibernation period for this species. The results of the present study emphasise the importance of forest and diverse residential areas including forest patches for the hedgehog’s nesting habitat.     

Nesting behaviour and foraging characteristics of Andrena cineraria (Hymenoptera: Andrenidae)

The current studies were carried out in the three experimental locations of Kashmir valley during 2013 to 2016. The species Andrena cineraria formed the dense nest aggregations in plan grounds, barren lands and hilly areas near the fruit orchards and other landscapes with clay loam soil type. The species start flying and foraging in the orchards from April till July. The nests were allodalous, 29–36 cm in depth, with cells located obliquely around the main barrow. The nests were dense with a maximum density of 11.09 nests/m² observed in landscapes of Budgam. The barrow diameters were found varying with depth from main entrance. The maximum barrow diameter recorded was 2.05 mm. At certain depth, the female constructs the first cell and the upper nest burrow is vertical and lower is oblique. The nest entrance is generally hidden under the tumulus. In the depth of average 30.48 cm, each cell directly opens to main burrow either alternately or unilaterally. The cell number, diameter, and length varied with depth. Foraging behaviour of A. cineraria on various fruit crops and other shrubs and social forestry trees were determined and the abundance, visitation rate, total visits and time spend per flower were found significant, especially on fruit crops. The significance of the studies is important for the melittologists, as it will help in the conservation of bee fauna. The study is also important in using this species for pollination purpose and would also help to detect and understand the possible pre-adaptation of species in temperate region of Kashmir valley.     

Termitarium volume as a determinant of invasion by obligatory termitophiles and inquilines in the nests of Constrictotermes cyphergaster (Termitidae, Nasutitermitinae)

A range of organisms can be found inside termite nests where the degree of association can vary from facultative to obligatory dependence. Studies of the dynamics of nest invasion are still unresolved, so how and when cohabitants enter termite nests remain open questions. This study analyzed one specific aspect of the dynamics of termite nest invasion by obligatory termitophiles and inquilines, i.e., whether cohabitants were more likely to invade a nest when it reached a critical nest size. We collected 36 Constrictotermes cyphergaster nests of different sizes and sampled their cohabitant fauna. Our results indicated that the invasion of C. cyphergaster nests by obligatory termitophiles and inquilines was dependent on nest size. There appeared to be a critical nest size above which nests were more prone to invasion. Above this size, there was a significantly higher likelihood of finding obligatory cohabitants. Termitophile species were observed in nests ≥2.2?L, whereas inquiline species were only occur in nests ≥13.6?L. This may indicate that the obligatory cohabitants studied here did not occupy C. cyphergaster nests at random and that they were dependent on features that made these nests suitable for cohabitation, which are linked to colony development.     

Addition of arthropod cocoons to house wren nests is correlated with delayed pairing

Males in the cavity-nesting house wren (Troglodytes aedon) frequentlyadd arthropod cocoons to their nests during building, possiblyas an ornamental cue for female choice. We tested this hypothesisby comparing the time to pairing for males that did and didnot add cocoons to their nests and for males in whose nestswe manipulated the number of cocoons prior to pairing. We alsotested the hypothesis that females acquire fitness-related benefitsby selecting mates based on their use of cocoons. The use ofcocoons by males was not consistently related to habitat, butthe number of cocoons added per nest increased during the courseof the breeding season. Contrary to prediction, the time topairing for males adding cocoons was significantly longer thanthat for males without cocoons in their nests at both unmanipulatedand experimental nests. There was also no consistent fitness-relatedbenefit for females related to the use of cocoons by their mates.Therefore, we conclude that females did not prefer males thatadded cocoons to their nests, and that the increased time topairing for males that add cocoons likely results in fitness-relatedcosts brought about by delayed breeding. Nonetheless, male housewrens routinely use cocoons, and why they do so remains unknown.     

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.     

Osmia cornifrons vitellogenin: cDNA cloning,structural analysis and developmental expression

Osmia cornifrons plays a major role in the pollination of orchards, but basic information on vitellogenin and oocyte development is limited. To better understand vitellogenin in hymenopteran insects, we cloned a cDNA encoding vitellogenin from the hornfaced bee O. cornifrons. Osmia cornifrons vitellogenin cDNA contains 5477 bp with an open reading frame of 1783 amino acid residues, and has a predicted molecular mass of approximately 200.21 kDa and a pI of 6.55. Osmia cornifrons vitellogenin possesses four consensus (RXXR/S) cleavage sites and has conserved DGXR and GL/ICG motifs in the C‐terminus. The deduced amino acid sequence of the O. cornifrons vitellogenin cDNA showed a 66% identity with Megachile rotundata, 53% to Apis mellifera, 51% to Bombus ignitus and 42%–30% with other hymenopteran insect vitellogenins. Phylogenetic analysis showed that O. cornifrons vitellogenin clustered with vitellogenins from Megachilidae, Apidae, Vespidae and Formicidae species but not with those from Pteromalidae, Aphelinidae or Ichneumonidae species. The expression profile of O. cornifrons vitellogenin mRNA during development revealed that O. cornifrons vitellogenin was first detected in the pupal stage and was continuously detected during the adult stage. Interestingly, O. cornifrons vitellogenin mRNA expression was low in mid‐diapause, then gradually increased beginning on day 3 of the newly emerged adult stage, and subsequently declined. These results suggest that the expression level of O. cornifrons vitellogenin mRNA is stage‐specific.     

Trap-nests used by Centris (Heterocentris) terminata Smith (Hymenoptera: Apidae, Centridini) at secondary Atlantic Forest fragments, in Salvador, Bahia State

Ninety-five nests of Centris (Heterocentris) terminata Smith were collected in trap-nests, during November/2001 and January/2003, at two fragments (PZGV e CFO-UFBA) of secondary Atlantic Forest, in Salvador, Bahia State (13 masculine01 W e 38 masculine30 S). The highest nest frequencies occurred from December to February (summer), with no nests foundations from August to October (winter - early spring). Two-hundred eight adults emerged from 347 brood cells, being 164 males and 116 females (1: 0.42). During the study period sex ratio was male biased (c2 = 9.342; gl = 10; P < 0.05). C. terminata nested in holes with diameters 6, 8, 10 mm, but 84,2% were constructed in 8 and 10 mm. nests had one to seven cells arranged in a linear series with the cells partitions built with a mixture of sand and resin or oil. Male is significantly smaller than female, which emerges from the first cells constructed. Immature mortality occurred in 14.1% of brood cells (n = 49), of which 13.0% were due fail in development and 1.2% due to parasitism of Coelioxys sp. (Hymenoptera: Megachilidae) e Tetraonyx sp. (Coleoptera: Meloidae). In the study site, weather, mainly pluviosity, rather than natural enemies influenced seasonal population abundance. The long period of nesting activity, local abundance and usage of trap nests, suggest the potential of C. terminata for management aiming at pollination of native and cultivated plants.     

Pre-ovipositional maternal care alleviates food stress of offspring in the flower beetle <Emphasis Type="Italic">Dicronocephalus wallichii</Emphasis>

Unlike most other flower beetles, females of Dicronocephalus wallichii exhibit nesting behaviour. The female constructs a burrow in the soil, cuts dead plant leaves into small pieces to provision the nest, and then lays one egg inside the nest. Hatched larvae have been thought to feed on the nest materials prepared by their mothers, but the effects of pre-ovipositional care on larval performance have not been tested. The hatched larvae were found to stay in the nest for 15–30 days until they consumed the nest materials. We examined whether the presence of provisioned nests enhanced larval performance under both benign and food-stress conditions. With high-nutrient soil, larval survival rate and growth speed were not affected by the presence of provisioned nests. By contrast, with low-nutrient soil, mortality of the larvae was much higher in the absence than in the presence of provisioned nests. The growth speed of larvae with nests located in low-nutrient soil was as high as those reared in high-nutrient soil. These results indicate that females alleviate the food stress of larvae during their initial developmental stage by constructing provisioned nests.