Nesting biology of four species of Trypoxylon (Trypargilum) (Hymenoptera: Crabronidae) in Chaco Serrano woodland, Central Argentina

Trypoxylon (Trypargilum) (Crabronidae) wasps are solitary spider predators that can build their nests in artificial trap-nests, which enables study of their nesting architecture and biology. Twenty traps (each containing 15-30 internodes of cane) were placed in each of nine sites of Chaco Serrano Woodland in Central Argentina (Córdoba) in October 2005, and were recovered in June 2006. We obtained 91 nests of four species of Trypoxylon (Trypargilum). In the laboratory, each Trypoxylon nest was sectioned longitudinally to study its architecture. The number of brood cells was counted, and the occurrence and length of vestibular and intercalary cells and the pre-closing plug space were recorded. We measured the diameter of the cane entrance, total length of the nest, length of each brood cell, maximum thickness of mud partitions and closing plug thickness. We also recorded the cell contents: the wasps, their natural enemies and the prey spiders. Mortality was assessed and the sex ratio calculated for each species. Finally, the nests were examined to help clarify the function of the vestibular cell. The nest architecture was similar in the four species, with linear brood cells located one after the other separated by mud partitions, as in other species of the subgenus Trypargilum. Forty-eight percent of the nests had vestibular cells, but only two had intercalary cells. The thickness of the mud partitions and the length of the brood cells differed among species and were related to the size of the emerged wasp. The diameter of the nest entrance was directly related to the average length of the fore-tibia. Sex ratios of all species did not deviate from 0.5. Mortality due to parasitoids (Eulophidae; Melittobia sp.) was similar among species, while the mortality due to cuckoo wasps (Chrysididae) in T. lactitarse was higher than in the other species. The presence of vestibular cells was not related either to the mortality due to natural enemies or to the orientation of the trap in the field. Spiders in the family Araneidae were the most frequently collected prey.     

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.     

Notes on the prey,nesting behaviour and natural enemies of three Bembix sand wasps (Hymenoptera: Crabronidae) in the Iberian Peninsula

Bembix is a cosmopolitan genus with more than 300 species distributed all over the world. The females of these wasps dig burrows in the ground and hunt for flies to feed their larvae. Current knowledge of the nesting behaviour of most Palaearctic species (around 50) is limited. In this study, we provide data on the nest structure, nesting activity, natural enemies and prey captured by three Bembix sand wasps present on the Iberian Peninsula: B. zonata, B. flavescens bolivari and B. merceti. We also analyze the prey spectrum of the three species, comparing our data with those available from previous studies. B. zonata exhibits the greatest selectivity index (and the narrowest niche width), capturing mainly Syrphidae to provision its nests. B. merceti shows the greatest niche width in comparison with the other two species. The inter-annual differences in the diet of B. merceti are important and are probably due to the ability of this wasp to change from one prey resource to another, depending on its availability.     

Opportunistic predation and offspring sex ratios of cicada-killer wasps (Sphecius speciosus Drury)

Abstract.  1. The way in which hunting of prey affects the sex ratio of the predator's offspring is not well understood. Female cicada-killer wasps are convenient for study because they specialise in capturing cicadas to provision their offspring. Cicada prey are nearly twice as heavy as the wasps that carry them, hence some degree of prey selectivity by the wasps is to be expected. It has been suggested that wasps bias their offspring towards females by foraging selectively for female prey, whereas there is some evidence that sex ratios are actually male-biased. This study was designed to establish the connection between foraging and offspring sex ratio.
2. Three, non-exclusive, hypotheses of selective predation were tested. The frequency of predation on different classes of prey in conjunction with their availability was estimated by intercepting the wasps on their way to their nests and by sampling cicadas in the environment. The hypothesis of selective predation was not supported; predation appears to be opportunistic and non-selective. Cicada prey weight was not a simple linear function of wasp weight, although the smallest wasps were constrained to carry small prey.
3. Wasp offspring (larvae) were excavated from subterranean nests and found to be male-biased (3 : 1 or 4 : 1) in 2 years. The observed ratios are close to expectation from Fisher's equal-investment model, taking account of sexual size dimorphism, and are evidently unrelated to the sex of the prey. A simple binomial probability rule of sex allocation provides a behavioural mechanism for producing the observed sex ratio of offspring.     

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.     

Abundance of wasps and prey consumption of paper wasps (Hymenoptera, Vespidae : Polistinae) in Northland, New Zealand

Polistine and vespine wasps were captured in Malaise traps in two fire-modified shrubland habitats of varying canopy height and composition at Lake Ohia, Northland, New Zealand. Prey consumption rates were calculated for the Asian paper wasp (Polistes chinensis antennalis) occupying these two areas of shrubland and a home garden in Whangarei, Northland. The sites were systematically searched for nests and wasp prey determined by intercepting foragers returning to nests. The Asian paper wasp predominated in the Malaise trap samples from the low- growing habitat while the German wasp (Vespula germanica) was more common in the taller vegetation type. The Asian paper wasp was more abundant than the German wasp in the samples in February and early March. Only four Australian paper wasps (Polistes humilis) and no common wasps (Vespula vulgaris) were caught. Asian paper wasps collected an estimated 15 000 prey loads per ha per season from one of the shrubland areas, and 478 000 prey loads per ha per season from the second area. These convert to estimates of 31 and 957 g per ha per season of invertebrate biomass removed by paper wasps from each habitat, respectively. The estimate for the garden site was 79 g per ha per season. Wasp nest densities varied between 20 and 210 nests per hectare. The biomass estimates are similar to average figures calculated for vespine wasps in scrubland and pasture. Both Asian paper wasps and Australian paper wasps preyed mainly on lepidopteran larvae. The cabbage white butterfly (Pieris rapae) was the most commonly collected species. Noctuid species were also well represented. Both male and female Asian paper wasps collected nectar in late March and early April.     

Hunting behaviour,prey capture and ant avoidance in the tropical social waspPolybia sevicea (Hymenoptera: Vespidae)

Summary Polybia sericea (Hymenoptera: Vespidae) prey foraging was studied by following individual foragers as they hunted in the field, by observing how wasps handled prey once they had captured it, and by observing wasps as they returned to the nest with prey. Wasps were most likely to forage for prey between 0700 and 1300 hours and between 1600 and 1700 hours. The prey foraging sequence consisted of the behaviours high flight, search, touch, land, groom, walk, bite and malaxate. Captured small prey were malaxated and carried to the nest. Wasps removed the gut from large prey and dragged the meat up a twig or grass stem. A load of the meat was then bitten off and malaxated; the remainder was cached while the wasp made an orientation flight and returned to the nest. The forager returned within minutes for the remainder of the prey. Experiments demonstrated that caching the prey remains above the ground rather than close to the ground, where the prey are generally captured, reduces the chance that the prey will be found and expropriated by ants.     

A NEST SHARED BY THE SOLITARY WASPS, CERCERIS ANTIPODES SMITH AND C. AUSTRALIS SAUSSURE (HYMENOPTERA: SPHECIDAE)

Abstract
Interspecific nest sharing between 2 species of sphecid wasps is recorded for the first time. A nest of Cerceris antipodes Smith appeared to have been joined by a female of C. australis Saussure. The nest contained 2 cells with a mixture of prey of both species, 26 older cells with C. antipodes prey and 6 younger cells with C. australis prey.     

The behaviour of Philanthus crabroniformis (Hymenoptera: Sphecidae)

Observations were made over a two year period on the nesting and hunting behaviour of a population of Philanthus crabroniformis Smith. Data are presented on the time and speed of provisioning, the capture of prey and the species taken, the design and location of nests, the approach to the nest with prey, and aggressive interactions between females. P. gibbosus Fabr., also nested in the same area and took much the same prey. Unlike other sympatric pairs of Philanthus which have been studied, the two species in question were not totally segregated by separate nesting seasons nor by mutually exclusive habitat preferences. One aspect of the behaviour of the two Philanthus which was highly distinctive was the manner in which prey-laden females approached their nests. Selection for divergence in approach patterns may have been exerted by miltogrammine flies, a major parasite of many digger wasps.     

Nesting habits of two spider wasps: Anoplius infuscatus and Episyron sp. (Hymenoptera: Pompilidae), with a review of the literature

Many behavioural aspects of Anoplius infuscatus (Vander Linden) and Episyron sp. Schioedte were examined in detail, both on a qualitative and, when possible, on a quantitative level, during a long-term field study in Northern Italy. Both wasps dig unicellular nests on bare soil, which are filled with a single spider to feed the brood. The main differences regard the way to hunt their prey, that of burrowing, that of transporting and introducing the spider into the nest, the habit of amputating spider legs and the abdominal movements during nest closing. On the whole, starting from prey capture to nest closure, A. infuscatus is significantly slower than Episyron. The observed ethological differences are well fitted to the exploitation of the different preys consisting in wolf spiders (Lycosidae) and orb weaving spiders (Araneidae), respectively. Some flexibility in the observed behaviour is described. A critical re-examination of the literature shows that, even if most results agree with those of previous authors, two controversial points remain open regarding Episyron species: the position of the egg (on the prey or on the cell wall) and the way to grasp the spider during tranport and introdution into the nest.     

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.     

Specialisation in prey capture drives coexistence among sympatric spider‐hunting wasps

1. Sister taxa that coexist in the same space and time often face competition due to the use of similar resources. However, some closely related species can adopt fine‐grained specialisation in resource use to coexist. This study investigated niche overlap between three sympatric spider‐hunting wasp species of the genus Trypoxylon (Hymenoptera: Crabronidae) known to nest in three of the habitats found in the study area. 2. First, the co‐occurrence of these wasp species in the three habitats was estimated, as a proxy for potential competition. Then, the following hypotheses were tested: (i) niche partitioning is seen more often between species that co‐occur in a habitat, whereas there is niche overlap between species nesting in distinct habitats (prey specialisation hypothesis); and (ii) wasp species capture prey according to their size (physical constraint hypothesis). 3. Two pairs of wasp species were found consistently nesting in the same habitat. Niche partitioning based on prey taxa occurred regardless of the habitat preference. It was also found that differences in the size of wasps reflected distinctions in the size of their prey. 4. These findings were consistent over the years, showing that the significance of specialisation in foraging activities and physical constraints during prey capture can play key roles in the coexistence of sympatric species. The distinctions in the foraging strategies of these wasps are discussed, as well as potential mechanisms driving the evolution in prey specialisation, with insights for future studies.     

中国南部狭腹胡蜂的生物学与地理分布(膜翅目：胡蜂总科)

1989-1991年与日本京都大学合作,对我国南部狭腹胡蜂区系、生态、行为学等方面进行了研究。本文主要报道狭腹胡蜂的生物学与地理分布。研究结果表明狭腹胡蜂的栖息和巢室建筑分野外和住宅两种环境。密侧狭腹胡蜂Parischnogaster mellyi(Sau.)的蜂巢密度在30-100m²之间的住户内,平均为8.25巢,最高“巢,最低为2巢。巢室材料分植物纤维和土两类。在东径99‘15'一104°98',北纬20°06'-22°55'之间范围内,除分布在海拔1000m以上类群外,一年四季都可进行活动,无越冬现象。分布范围最北限止于贵州东北部。     

Morphological distance and inter-nest distance account for intra-specific prey overlap in digger wasps (Hymenoptera: Crabronidae)

Although inter-individual diet variation is common in predatory wasp populations, the factors accounting for such variation are still largely unknown. Here, we asked if paired diet dissimilarity in three species of digger wasps correlates with morphological distance and inter-nest distance, two factors previously linked to diet partitioning in vertebrates. Results sharply differed among species and generations. All sampled populations showed significant inter-individual diet variation for prey taxa, but only in half of the cases for prey size. In one generation of two species [Bembix zonata Klug and Stizus continuus (Klug)], similar-sized wasps had similar prey taxonomic spectra (and for S. continuus also similar prey size spectra), a phenomenon which probably reduces intra-specific competition. In addition, B. zonata females nesting closer to each other had more similar prey taxonomic spectra, suggesting that distant females probably hunt on different patches that harbour different prey species. For the females of a further species (Bembix merceti Parker), pairwise size difference and inter-nest distance did not affect prey dissimilarity. Both morphological distance and inter-nest distance are potentially important in shaping the overlap of individual resource use in wasps, though probably only in certain conditions such as a highly clumped distribution of nests and size-related constraints on prey selection.     

Patterns of nest provisioning and parental investment in the solitary digger wasp Ammophila sabulosa

Abstract.

• 1 In nest-building wasps there is a continuum from species that provision each offspring cell with one large prey item (single provisioning) to species that provision each cell with many small items (multiple provisioning). The significance of number of prey per cell was examined during two seasons in Ammophila sabulosa (L.), a species that provisions between one and five caterpillars per cell.
• 2 There was no difference between the total weight of prey in singly-provisioned and multiply-provisioned cells: individual caterpillars placed in multiply-provisioned cells were smaller.
• 3 Small caterpillars were captured and transported to the nest faster than large ones.
• 4 If the first prey provisioned was large, the cell was usually then permanently closed. If the first prey was small, additional prey were captured, and females appeared to become more selective: additional prey were rarely large.
• 5 It is suggested that the flexible provisioning strategy used by A.sabulosa allows a wide range of prey sizes to be utilized without affecting maternal control of offspring size and sex. This could be important when available prey sizes vary temporally and spatially.
• 6 There was a male-biased first generation investment ratio. Female-producing cells were provisioned with a higher total prey weight than male-producing cells, and conversion of prey weight to adult weight was more efficient for females.
• 7 Total prey weight provisioned is probably a good indirect measure of parental time investment in A.sabulosa. Other measures, particularly number of prey provisioned, will be less accurate.

     

Effects of wax bloom variation in Brassica oleracea on foraging by a vespid wasp

Foraging by individual Polistes dominulus (Christ) (Hymenoptera: Vespidae) was observed and compared on Brassica oleracea L. plants that differed in surface wax bloom. Twenty-six wasps, previously trained to forage on plants for 4th instar Trichoplusia ni (Walker), were presented in the greenhouse with mixtures of plants with normal wax bloom or genetically reduced wax bloom, and on which T. ni caterpillars had been placed. During foraging, the wasps were observed to slip significantly more frequently from the leaf surfaces of normal-wax bloom plants than from reduced-wax bloom plants (129 vs 63 occurrences) and to retrieve significantly fewer pieces of caterpillar after attack on normal-wax bloom plants (151 vs 223 pieces). Altogether the wasps retrieved 333 caterpillar pieces from reduced-wax bloom plants and 248 pieces from normal-wax bloom plants. Despite these differences, the number of caterpillars attacked and killed did not differ between the two wax bloom types (116 vs 121), nor did handling time for individual attacks (time from contact with prey until prey piece was carried to the nest) (170±12.5 s vs 180±10.5 s). Thus, in contrast with previous reports for smaller predators, wax bloom variation in B. oleracea did not influence the effectiveness of P. dominulus as a predator of T. ni.     

Nutrient Dynamics of a Swarm-founding Social Wasp Species,Polybia occidentalis (Hymenoptera: Vespidae)

The kinds, rates of acquisition, inter-individual transfers, and intra-colonial movements of nutrients were ascertained for the advanced eusocial paper wasp Polybia occidentalis (Olivier). Foraging worker wasps (“foragers”) bring arthropod prey and nutritive liquids (“nectar”) to the nest, and these are usually transferred to nest workers (“receivers”) on the outer nest envelope. Arthropod prey items, which are brought intact to the nest, are malaxated by one or more receivers before being fed to larvae; malaxating adults retain a portion of the hemolymph for their own nourishment. Nectar is usually transferred (via adult-adult torphallaxis) from foragers to receivers on the nest envelope; some nectar is given to larvae, and all adults that imbibe it retain at least some for their own nourishment. Larval saliva of P. occidentalis contains glucose, protein, and free amino acids and so is highly nutritive; the nutrient content of the saliva closely resembles that of the saliva of other social wasp taxa. Adult wasps imbibe larval saliva, but very little is apparently transferred by those adults (via trophallaxis) to nestmates. Brood cannibalism was infrequent during this study. Adult worker and male wasps possess chymotrypsin-like and trypsin-like enzyme in their midguts and so are likely capable of protein digestion. The midgut proteases show an age-correlated variation in concentration. Pulp foragers are significantly smaller and lighter in weight than are receivers.     

Comparative characteristics of nests of Vespula germanica (F.) and Vespula vulgaris (L.) (Hymenoptera: Vespinae) from Christchurch City,New Zealand

Abstract

From July 1987 to June 1989, 141 nests and 365 nests, respectively, of the adventive social vespines the German wasp, Vespula germanica (F.), and the common wasp V. vulgaris (L.), from the City of Christchurch, New Zealand, were evaluated for seven major characteristics.

There were few, if any, differences in nest sites, and from spring to early summer in nest traffic, nest size, and numbers of combs. However, Getman wasps showed no preference for direction of nest entrances, while common wasp nests were more numerous in areas most exposed to the morning sun. Some Getman wasp nests survived the winter and began producing new worker cells by late June, but all common wasp nests died by June. Because common wasp nests have been reported surviving the winter in beech forest, which produces honey dew, and honey dew is not available in Christchurch City, common wasp nest survival over winter may be more dependent upon carbohydrates and/or prey dependent upon carbohydrates than the survival of German wasp nests. The presence of large, expanding wasp nests from early spring must impose localised predation pressures virtually unknown in the Northern Hemisphere where overwintering nests are rare. Wasp population dynamics, and impacts of wasps on fauna, are likely to vary between different geographical areas of New Zealand, but as targets for biological control, the two wasp species can generally be considered to be quite similar.     

Predatory behaviour in the ant-like wasp Methocha stygia (Say) (Hymenoptera: Tiphiidae)

An examination of prey-orienting behaviour of Methocha stygia has demonstrated that adult females regularly transport prey (Cicindela larvae) when they are removed from their burrows. On a single occasion a wasp was also observed to utilize a naturally occurring depression in the soil as a nest site, instead of depending on the prey's own burrow. This flexibility in response might be a carryover of the behaviour that surely existed in the ancestors of the methochine wasps, insects which were close to if not identical with the ancestors of the ants. When Methocha females were crowded together with prey in the laboratory, frequent avoidance and aggressive behaviours were noted, but no sign of any form of co-operation.     

Elevational Patterns of Diversity and Abundance of Eusocial Paper Wasps (Vespidae) in Costa Rica

We used a standard sampling protocol to measure elevational patterns of species richness and abundance of eusocial paper wasps (Hymenoptera: Vespidae) in Costa Rica. The sample transect of six sites spanned approximately 2000 m in elevation from lowland to montane forest. Species accumulation curves and species richness estimates both document a low elevation peak in paper wasp species richness at 50 and 300 m asl, with a decline in species richness at higher elevations. Comparison of species composition among elevations revealed strong species turnover from a rich lowland fauna to a depauperate, but distinct, montane fauna. We also observed a general trend toward a greater abundance of paper wasps at higher elevations, a pattern not usually observed in eusocial insects. Army ant species that prey on paper wasps declined in abundance with elevation across the sample transect, a pattern that has been observed at other sites. We discuss the possibility that elevational changes in predation pressure affect variation in paper wasp abundance and species richness. Eusocial paper wasp species employ one of two modes of colony founding, independent and swarm founding. We found that the total abundance of individual swarm-founding wasps was higher at all elevations than the abundance of independent-founding wasps, supporting previous suggestions that Neotropical swarm founders are more successful ecologically.