Biology of the solitary wasp Trypoxylon (Trypargilum) agamemnon Richards 1934 (Hymenoptera: Crabronidae) in trap‐nests

Buschini, M.L.T. and Fajardo, S. 2009. Biology of the solitary wasp Trypoxylon (Trypargilum) agamemnon Richards 1934 (Hymenoptera: Crabronidae) in trap‐nests. —Acta Zoologica (Stockholm) 91 : 426–432. Some Trypoxylon species build their nests in preexisting tubular cavities like hollow stems and beetle borings in branches. Study of the biology of these insects is relatively easy because the females of these wasps nest with enormous success in trap‐nests. The aim of this study was to investigate the abundance, seasonality and life‐history of Trypoxylon agamemnon. For capture of these insects, trap‐nests were installed in the Parque Municipal das Araucárias in araucaria forest, grassland and swamp, from December, 2001 to December, 2005. Two hundred and ninety seven nests were obtained. They were constructed more often during the summer (from December to April). The nests were built only in araucaria forest and consisted of a linear series of cells, divided by mud partitions, whose number varied from 1 to 7. Normally they have only one vestibular cell. The inner cells had been provisioned, usually with spiders of Anyphaenidae family. Sex‐ratio was strongly female biased. Its main natural enemies included Chrysididae, Ichneumonidae and Tachinidae.     

Notes on the biology of Trypoxylon (Trypargilum) opacum Brèthes (Hymenoptera; Crabronidae) in Southern Brazil.

The present study investigated the abundance, seasonality and various life-history traits of Trypoxylon (Trypargilum) opacum. Using trap-nests, 320 nests of T. opacum were collected in the Parque Municipal das Araucárias in Southern Brazil (25 degrees 23' 36" S and 51 degrees 27' 19" W) over a 3 year period. Nesting was more frequent during the warm season. Nests consisted of a linear series of 1 to 8 brood cells separated by mud partitions, usually followed by an empty vestibular cell and final-closure mud plug. Brood cells were most commonly provisioned with spiders of the family Araneidae. Sex-ratio was strongly female biased, 3.4:1 females:males. Natural enemies attacking nests T. opacum included chrysidids, ichneumonids, sarcophagids, bombyliids and ants.     

The Significance of the Vestibular Cell in Trap Nesting Wasps (Hymenoptera: Crabronidae): Does its Presence Reduce Mortality?

In a study carried out with the trap-nesting solitary wasp Trypoxylon attenuatum (Hymenoptera: Crabronidae) in 2000, we analysed the significance of different nest characteristics, like the vestibular cell and empty intercalary cells, and its role in the reduction of mortality. Analysis of the presence of the vestibular cell suggested that it represents a nest completion structure, whose presence reduces mortality in the initial stages (M1) but does not affect that due to chrysidid parasitoids (MP). The length of such vestibular cell was found to depend mainly on the space between the last provisioned cell and the exterior, pointing to a possible role in the reduction of the remaining space that could be used by other females to establish a nest exterior to the completed one. The empty intercalary cells were smaller than the vestibular cells, did not affect M1 or MP mortality rates in the nest, and exhibited an “aggregated” pattern of occurrence. This suggests that they could be the result of “abnormal” behaviour by the female owning the nest. The total mortality rate decreased with the increase in the number of cells in the nests, also being greater in nests established at low heights above the ground. M1 was lower in the innermost cells, while MP increased towards the interior and in the nests with more cells. The distribution in the nests of cells parasitized by Trichrysis cyanea (the main natural enemy found in the wasp population studied) reflects an “aggregated” pattern, suggesting repeated attacks against certain nests.     

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.     

Comparative aspects of the biology of five Auplopus species (Hymenoptera; Pompilidae; Pepsinae) from Brazil

This study was carried out at the Parque Municipal das Araucárias, in the municipality of Guarapuava, state of Paraná, southern Brazil, from December 2001 to December 2004. Sixty nests belonging to five Auplopus species were collected using trap-nests. The nest cells of all species were built with mud and shaped like a barrel or cylinder, mostly in linear arrangement. Differences between species lie in the colour of the mud and the way in which they organize the mud pellets during the construction of the cells. Few vestibular and intercalary cells were built. The number of cells varied according to the species, never surpassing six cells per nest. We observed a great variation between the five species related to sex ratio and the number of generations per year. Some species presented a delayed life cycle with juveniles entering diapause, others presented an additional short life cycle with no diapause.     

Nesting and biology of Jucancistrocerus caspicus (hymenoptera,vespidae, eumeninae)

Females of Jucancistrocerus caspicus nest in dense clay ground on the vertical surface of cliffs. The nests contain 1–9 cells (on average 2.8) and have a linear-branched construction. Females surmount the entrance of the burrow with a curved chimney which has a laced structure. The cells are positioned vertically in the main burrow and obliquely or horizontally in the lateral tunnels; the cells in a row are separated with double partitions. The size of the cells is 7–9 × 4–4.5 mm, the diameter of the nest burrow is 4 mm. The egg is laid before provisioning and is attached to the cell ceiling with a filament. Females hunt for weevil larvae and store 23–33 larvae (on average 27.8) in each cell. The species is univoltine, with prepupae hibernating in their cocoons. The nests are parasitized by the cuckoo wasps Chrysis rutilans which cause 11.5% of brood mortality. Adult wasps are killed by the spiders Pholcus sp. living near the nests.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Nesting biology of Pareumenes quadrispinosus (de Saussure, 1855) (Hymenoptera: Vespidae: Eumeninae) in trap nests in North Vietnam

Nests of Pareumenes quadrispinosus (de Saussure, 1855) were obtained at Me Linh Station for Biodiversity (323 nests) and Tam Dao Town (283 nests), Vinh Phuc Province, as well as at Phu Luong, Thai Nguyen Province (9 nests) and Kim Boi, Hoa Binh Province (62 nests). The wasps nested in segments of bamboo canes and reed stems, 5 to 18 mm in internal diameter. Each nest consisted of a linear series of one to four cells, separated by mud partitions. Brood cells were provisioned with caterpillars and eggs were attached to the ceiling of the cells by thin threads. The life history and sex ratio of this species nesting in trap nests in North Vietnam were recorded from mid-April to last October. The species is multivoltine, with likely up to four generations per year. It has two alternative life histories (diapause and direct development) and overlapping generations. The sex ratio is strongly female-biased. Only 46% of the provisioned cells were successful; the others were damaged by nine parasitoid species or died during development for unknown reasons.     

Nesting behavior of Podium denticulatum Smith (Hymenoptera: Sphecidae)

The nesting behavior of Podium denticulatum Smith was studied on the campus of Ribeir?o Preto of the Universidade de S?o Paulo, SP, Brazil, from September 2003 to August 2005. The wasps established their nests in bamboo canes ranging from 11.4 cm to 26.2 cm in length and from 0.7 cm to 1.8 cm in internal diameter. Podium denticulatum nested almost exclusively in the hot and wet season (September-April), producing at least five generations per year. The cell provisioning was made with adult and nymphal cockroaches (Blattellidae) which were arranged venter-up and with the head inward toward the inner end of the cell. The construction of a temporary closure occurred in cells that took more than one day to be provisioned. The cells provisioned with a greater number of prey were more likely to produce females than males. The nests included 1-6 brood cells separated by mud partitions and arranged in a linear series. The innermost cells of the nests produced females, and the outermost cells produced males. Nests were parasitized by Eulophidae (Melittobia sp.), Chrysididae and Tachinidae.     

Nesting biology of the potter wasp Rhynchium brunneum brunneum (Fabricius, 1793) (Hymenoptera: Vespidae: Eumeninae) in North Vietnam

Rhynchium brunneum brunneum (Fabricius, 1793) is a common species using trap nests in North Vietnam. The females chose the nest traps with diameters ranging from 5.5 to 17 mm. Nests consisted of a linear series of one to eleven brood cells separated by mud partitions. Brood cells were provisioned with caterpillars, and eggs were attached to the ceiling of the cells by a thin filament. The life history and sex ratio data of this species were recorded from April to early November. Its sex ratio is strongly male-biased, being multivoltine, likely with four generations per year, the last one overwintering in the prepupal stage. Nesting activity of the species was described with major activities such as nesting site selection, oviposition, prey collecting, and applying cell material. Only 53.3% of the provisioned cells were successful; the others were damaged by six parasitoid species or died during development for unknown reasons.     

Intraspecific parasitism and nest defence in the solitary pompilid wasp Anoplius viaticus (Hymenoptera: Pompilidae)

The nesting behaviour of individually marked female pompilid wasps, Anoplius viaticus , was observed at a Breckland heath site with particular emphasis on intraspecific parasitism and nest defence. Prey was stolen from conspecifics while it was being carried to the nest site, and while it was left unattended during nest construction. Females also appeared to brood parasitize each other's completed nests. Parasitism appeared to be opportunistic. Brood parasitism may be a tactic by which time-limited females can increase their fecundity. By placing prey in vegetation tufts during nest construction, females may reduce the risk of prey theft. An individual female's successive unicellular nests were clustered and therefore easier to defend, in many ways resembling a multicellular nest. Females defended their clusters vigorously, visiting them every few minutes during foraging and expelling conspecifics from the vicinity. This type of nest defence may be costly, and has rarely been observed in solitary wasps.     

Patterns of Nest Occupancy and Provisioning in Cerceris rufopicta Smith (Hymenoptera: Sphecidae)

Data are presented on prey choice, nest architecture, natural enemies, daily activity patterns, and seasonal patterns of nest occupancy in a population of Cerceris rufopicta Smith in northeastern Kansas. Most behaviors observed in this population are quite similar to those reported from populations in Florida, North Carolina, Pennsylvania, and New York. As in the eastern populations, by far the most common prey are two species of beetles in the subfamily Eumolpinae (Chrysomelidae), but late in the season at least one wasp began provisioning primarily with Eugnamptus angustatus. (Herbst), a weevil in the family Rhynchitidae, not previously recorded as prey of C. rufopicta. Nest architecture and natural enemies are very similar to what has been reported for the eastern populations, but provisioning activity in the Kansas population occupies a much greater part of the day, with some wasps continuing to provision even after sunset. This difference is not because individual wasps in Kansas are active for a longer period of time, but because the activity schedules of different individuals within the Kansas aggregation are not closely synchronized. Approximately half of the nests that were monitored over a period of three and a half weeks were occupied by a single wasp. Other nests changed ownership during the course of the study, either because the original owner disappeared and another wasp moved in, or because one wasp drove another away from a nest. During the time that ownership of a nest was being contested it might be simultaneously used by more than one wasp, but no evidence of cooperative joint nesting was seen in this population.     

Nesting biology of Trypoxylon (Trypargilum) lactitarse Saussure (Hymenoptera, Crabronidae) in trap-nests in Southern Brazil.

This study was carried in the Parque Municipal das Araucárias in the municipality of Guarapuava, state of Paraná, Southern Brazil. Three hundred and sixty five nests of T. lactitarse were obtained using trap-nests of 0.7, 1.0, and 1.3 cm in diameter. All of them had similar architecture, regardless of the diameter of the trap-nest. Completed nests consisted of a linear series of brood cells whose average number per nest was of 3.3, 4.0 and 3.6 for the nests with 0.7 cm, 1.0 cm and 1.3 cm in diameter, respectively. They were constructed more often during the summer. T. lactitarse had two types of life cycles: direct development (without diapause), and delayed development (with diapause during winter). Natural enemies included Chrysididae, Sarcophagidae, Dolichopodidae and Ichneumonidae. Out of 1,353 identified spider prey, 1,313 belonged to the Araneidae family.     

The comparative distribution and population dynamics in Trinidad of Sceliphron fistularium (Dahlbom) and S. asiaticum (L.) (Hymenoptera: Sphecidae)

Two sphecid wasps, Sceliphron asiaticum (L.) and S. fistularium (Dahlbom), which build mud nests and capture spider prey, had overlapping but distinctly different distributions in Trinidad. Sceliphron asiaticum was associated with drier (<2000 mm mean annual precipitation) and less forested areas. The analysis of existing data on these species revealed a similar differential distribution throughout the neotropics. Based on the dissection of their old nests collected at 42 sites in Trinidad, retrospective life tables were constructed and estimates of fecundity made. Sceliphron asiaticum tended to form denser nesting associations and, while most of the common parasitoids were shared, had overall higher developmental mortality. Although such mortality was density-dependent in both species, where the two wasps nested together mortality from Melittobia (Eulophidae, Chalcidoidea) was higher in S. asiaticum . Both species were protarrhenotokous and had similar field fecundities (7–8/♀), but the rate of oviposition may be greater in S. asiaticum since far less mud was used in the construction of its cells. Both species had >50° mortality of the females between emergence and starting to nest. The regulation of numbers of these wasps in relation to that of S. assimile (Dahlbom) in Jamaica is discussed.     

Comparative analysis of thermoregulation in the nests of certain species of social insects

Studies have been made on thermal regulation in the nests of families of the honey bee Apis mellifera, wasp Dolihovespula silvestris and bumblebees Bombus terrestris, B. agrorum and B. lapidaris during their maximum development. It was shown that thermoregulation significantly stimulated the brood in the nest. Among the species investigated, the highest thermoregulatory capacities are exhibited by honey bees, this fact being associated with the large number of individuals in their colonies. During cooling, bees group around the brood and their bodies make a cover of thermal insulation. Sheltering of the nests plays the main role in heat preservation of wasps and bumblebees. The latter, using contact method of heating of cells with the brood, provide rather constant temperature for its development. Honey bees, wasps and bumblebees react to overheating in the nests essentially in a similar way, i. e. by active aeration of their nests by vigorous wing beatings. The frequency of beatings increases with the increase in temperature.     

A Unique Nest-Protection Strategy in a New Species of Spider Wasp

Hymenoptera show a great variation in reproductive potential and nesting behavior, from thousands of eggs in sawflies to just a dozen in nest-provisioning wasps. Reduction in reproductive potential in evolutionary derived Hymenoptera is often facilitated by advanced behavioral mechanisms and nesting strategies. Here we describe a surprising nesting behavior that was previously unknown in the entire animal kingdom: the use of a vestibular cell filled with dead ants in a new spider wasp (Hymenoptera: Pompilidae) species collected with trap nests in South-East China. We scientifically describe the ‘Bone-house Wasp’ as Deuteragenia ossarium sp. nov., named after graveyard bone-houses or ossuaries. We show that D. ossarium nests are less vulnerable to natural enemies than nests of other sympatric trap-nesting wasps, suggesting an effective nest protection strategy, most likely by utilizing chemical cues emanating from the dead ants.     

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.     

Relationships between Nest Architecture and Behavior in <Emphasis Type="Italic">Xylocopa virginica</Emphasis> (Hymenoptera: Apidae)

The large carpenter bees nest in reeds, stems and wood. Many other Hymenoptera, including many wasps and some bees, have strong associations between the physical elements of their nests and behavior. Nests of the North American carpenter bee, Xylocopa virginica, were collected in southern Ontario. Nest architecture elements were examined with respect to their impact on life-history elements. In particular, it was determined that the brood cells are provisioned sequentially both within and among branches. There was also no detectable pattern of sex allocation in broods. Nests with branched architecture contained more foundresses, but these foundresses did not appear to contribute in the construction of larger nests. These findings are discussed with respect to other bees, and to the social structure of Xylocopa virginica.     

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.     

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.