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.     

BIRD-INSECT NESTING ASSOCIATIONS IN AUSTRALIA.

Australian records, beginning in 1866, of birds associating with aggressive insects for nesting purposes are assembled. Five species of the genus Gerygone are known to associate with wasps, more or less consistently, in tropical areas, and another species is reported as associating with wasps in the tropics and ants in temperate areas. Such birds deliberately seek the society of the insects, by which they are recognized and tolerated.
If protection is the motive, it would seem to be aimed against reptiles rather than against other birds or mammals. Other possible reasons for the practice ate suggested.
Australian records of insects frequenting birds' nests also are summarized; these include parasites such as the blood-sucking larvae of a fly, together with moth, beetle and fly larvae that serve a beneficial purpose as scavengers, and, in addition, wasps and beetles parasitic on the other insects. On the question of how the insects find the nests at appropriate times, it is suggested that some are guided by odour and others by some undetermined influence.     

西藏波密柑马蜂的筑巢习性（英文）

柑马蜂Polistes mandarinus在不同的环境中会利用植物材料制作各种不同的蜂巢。我们在2009年5-8月间观察研究了西藏自治区波密县的柑马蜂筑巢习性, 并收集62个蜂巢。柑马蜂利用植物纤维和树脂将它们的巢穴筑于当地居民的木屋檐下。收集的42个在用蜂巢中, 8月末平均拥有79.64±65.28个蜂房, 13.1±10.68个蜂蛹。20个弃用蜂巢中, 平均拥有12.95±3.99个蜂房。8月末, 成功的柑马蜂群拥有约80个个体。研究地的柑马蜂采取3种方式来筑巢, 但以第1种方式最为普遍。第1种筑巢方式筑造的蜂巢以两个蜂房连接巢柄, 结构最为稳定, 所以也应用最广, 后两种筑巢方式筑造的蜂巢均以一个蜂房连接巢柄, 是由于越冬雌蜂在找不到合适的筑巢材料时才采取的建巢方式。     

Nest site competition between cavity nesting passerines and golden paper wasps Polistes fuscatus

Nest boxes provide sheltered nesting sites for both passerines and paper wasps. Although neither wasps nor birds appear to evict the other once one is fully established, it is unclear which is the dominant competitor at the onset of the breeding season. Using wire mesh, we excluded birds but not golden paper wasps Polistesfuscatus from alternating boxes along a transect through edge habitat in North Carolina from 2006 – 2008. If wasps dominate Carolina chickadees Poecile carolinensis and eastern bluebirds Sialia sialis during the early spring (all have similar nest initiation dates), we would expect wasps to settle in both box types at equal frequencies. However, if birds dominate wasps, we would expect wasp nests to be concentrated in “bird‐proof” boxes. We found wasps in bird‐proof boxes significantly more often than in bird‐accessible boxes, indicating that secondary‐cavity nesting birds are able to exclude wasps from available nest sites. Additionally, we found that during the period of nest initiation, birds usurp wasps more often than vice versa.     

The control of nest depth in a digger wasp (Sphex ichneumoneus L.)

Golden digger wasps, Sphex ichneumoneus L. (Sphecidae), are a solitary, ground-nesting species that dig burrows to particular depths in the soil. I develop and evaluate alternative hypeses about the mechanisms controlling digging behaviour. By altering the wasps' burrows as thare digging, I show that they are not digging for some prescribed length of time nor are they diggingtil they reach some suitable environmental characteristic deep in the soil. Rather, they appear be digging until they reach a particular tunnel length, making corrections if the tunnel is too shallor too deep. This distance can be altered somewhat by surface environmental conditions.     

An overview of proximate factors affecting the nesting behavior of solitary wasps and bees (Hymenoptera: Aculeata) in preexisting cavities in wood

Guilds of Aculeate solitary wasps and bees that nest in preexisting cavities in wood are important components of terrestrial ecosystems because they engage in several ecological interactions (e.g. predation and pollination) with other species of plants and animals. Spatial and temporal variations in richness and abundance of solitary wasps and bees can be related to changes in environmental structure and in the diversity of other groups of organisms. The nesting period of these Aculeata is their most critical life cycle stage. Females of solitary wasp and bee species invest relatively more time constructing and provisioning their nests than do females of social species. Differently from species that nest in the soil or construct exposed nests, the main factors affecting the reproductive success of solitary species nesting in preexisting wood holes are still unknown. Our objective is to provide an overview of the role of proximate causes of nesting failure or success among solitary wasps and bees (Aculeata), for designing effective conservation and management strategies for these Hymenoptera.     

The effect of hind-tibial spurs on digging rate in female eastern cicada killers

Abstract.  1. The function of the large hind-tibial spurs of female eastern cicada killer wasps ( Sphecius speciosus ), thought to be adaptations to digging, was examined by measuring the effect of spurs on digging rate.
2. Spurs were removed from one group of wasps and left intact in control specimens.
3. Wasps with intact spurs removed soil from the burrow at a rate of 0.98 ± 0.11 (16) g dry mass min⁻¹ [mean ± SEM ( N )]. Removal of spurs results in an average digging rate just over one-half that, 0.54 ± 0.13 (9) g min⁻¹.
4. The difference in digging rate was primarily because of a twofold greater mass of soil unloaded by intact wasps each time they exited the burrow, rather than the rate at which such loads were delivered.
5. Such inefficiency results in an estimated additional 8.2 h requirement to fully excavate an average burrow, or a 1–4% reduction in time available for foraging in spur-ablated wasps.
6. Calculations suggest that the time lost to digging would result in a 3.9–19.5% reduction in foraging success for spurless females.
7. These temporal and resulting foraging advantages of hind-tibial spurs presumably provided selection pressure for their evolution from smaller, pre-existing setae.     

The biology of the eumenid mud-wasp Pachodynerus msidens in trapnests

Abstract. 1. The biology of the eumenid mud-wasp Pachodynems nasidens (Latr.) was studied in trapnests.
2. Females nested in traps placed in the shade and some nested in the same traps from which they had emerged.
3. The nesting behaviour and nest structure were generally similar to those of other eumenids that nest in borings, but a temporary crescent-shaped plug at the nest entrance and the use of both dry soil and mud for nest-building seems unique to P.nasidens .
4. Like most other eumenids, P.nasidens collected caterpillars, mainly Pyral-idae, Olethreutidae, Alucitidae and Thyrididae.
5. In linear nests, the innermost cells contained females and the outermost cells males, resulting in proterandry. One female can make several nests, each showing proterandry.
6. Female immatures were larger and took more time to develop than males. Also, their cells were larger and stocked with more food than male cells, hence needing more time to be provisioned.
7. The greater 'cost' to produce a female and a sex ratio biased toward females in short traps and about unity in longer traps, leads to a comment on parental investment.     

Long-maintained social–ecological systems and genetic diversity in coexisting insects: A case of the straw-thatched roof nesting solitary wasp Symmorphus apiciornatus (Hymenoptera: Vespidae: Eumeninae)

Thatched roofs are artificial environments that serve as nesting sites for a variety of insects, including tube-nesting wasps, but they have been declining drastically in recent years. In this study, we investigated the nesting habits of a eumenine wasp, Symmorphus apiciornatus, nesting in a group at thatched roofs in a northern part of Ibaraki Prefecture, central Japan. Genetic diversity and genetic structure in the area were also investigated. Sex ratios at most study sites were not biased. Males emerged first and waited for females to mate around the natal nests, suggesting a high possibility of breeding between the wasps having emerged at a given thatched roof under the recent situation where there has been a drastic reduction in the number of houses with a thatched roof. Nevertheless, the genetic diversity in a given local population or a thatched roof was relatively high and significant genetic differentiation among thatched roofs was not detected.     

Soil temperature, digging behaviour, and the adaptive value of nest depth in South American species of Acromyrmex leaf-cutting ants

In leaf-cutting ants, workers are expected to excavate the nest at a soil depth that provides suitable temperatures, since the symbiotic fungus cultivated inside nest chambers is highly dependent on temperature for proper growth. We hypothesize that the different nesting habits observed in Acromyrmex leaf-cutting ants in the South American continent, i.e. superficial and subterranean nests, depend on the occurrence, across the soil profile, of the temperature range preferred by workers for digging. To test this hypothesis, we first explored whether the nesting habits in the genus Acromyrmex are correlated with the prevailing soil temperature regimes at the reported nest locations. Second, we experimentally investigated whether Acromyrmex workers engaged in digging use soil temperature as a cue to decide where to excavate the nest. A bibliographic survey of nesting habits of 21 South American Acromyrmex species indicated that nesting habits are correlated with the soil temperature regimes: the warmer the soil at the nesting site, the higher the number of species inhabiting subterranean nests, as compared to superficial nests. For those species showing nesting plasticity, subterranean nests occurred in hot soils, and superficial nests in cold ones. Experimental results indicated that Acromyrmex lundi workers use soil temperature as an orientation cue to decide where to start digging, and respond to rising and falling soil temperatures by moving to alternative digging places, or by stopping digging, respectively. The soil temperature range preferred for digging, between 20°C and maximally 30.6°C, matched the range at which colony growth would be maximized. It is suggested that temperature-sensitive digging guides digging workers towards their preferred range of soil temperature. Workers’ thermopreferences lead to a concentration of digging activity at the soil layers where the preferred range occurs, and therefore, to the construction of superficial nests in cold soils, and subterranean ones in hot soils. The adaptive value of the temperature-related nesting habits, and the temperature-sensitive digging, is further discussed.     

Conditional female strategies influence hatching success in a communally nesting iguana

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

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.     

Normal and atypical nesting behaviour of Copris lunaris (L.): comparison with related species (Coleoptera, Scarabaeidae)

Abstract. 1. The lifecycle, mating and larval behaviour of Clunaris are described. Adults appeared in the autumn and nested in the following spring. The female beetle remained in the nest with the brood and could nest again the following year.
2. Nesting was initiated when virgin females were mated in the spring. Brood balls were formed by techniques sirnilar to those used by Scarabaeini. The female beetle left the nest soon after the first imagos broke out of the brood balls.
3. Nesting behaviour was readily modified by external conditions. Many parts of the sequence could be repeated or omitted. The female beetle left the nest if the brood was removed, but she remained for longer than usual if younger brood was substituted near the end of the normal nesting period.
4. Certain experimental conditions released behaviour patterns typical of other species. These were formation of superficial nests or of two-chambered nests, oviposition before completing the brood ball, and coating of the brood balls with soil (all found in other Coprina), as well as ball rolling and ball burial (found in Scarabaeini). The results are discussed in relation to the evolution of Copris nesting behaviour.     

Nest-site characteristics of the Red-knobbed Hornbill Aceros cassidix and Sulawesi Dwarf Hornbill Penelopides exarhatus

Nest-site characteristics of two hornbill species, the Red-knobbed Hornbill Aceros cassidix and the Sulawesi Dwarf Hornbill Penelopides exarhatus , are presented and compared at two sites in north Sulawesi, Indonesia. Seventy-four Red-knobbed Hornbill and 12 Sulawesi Dwarf Hornbill nests were located. Sulawesi Dwarf Hornbill nests were at lower densities and more evenly spaced than Red-knobbed Hornbill nests. Although selective, neither species appeared to be nest-site limited, except perhaps for the Red-knobbed Hornbill in hill forest. Principal components analysis indicated that Red-knobbed Hornbills were 'catholic' in their preferences, although nests were generally in tall, mature specimens of large girth, located high and in the upper half of the tree. The Sulawesi Dwarf Hornbill nested exclusively in live trees and nests were on average located half way up in tall canopy trees with large girths. Within certain parameters, both species were flexible with respect to their nest cavity size and shape requirements; Red-knobbed Hornbills choose sites with larger entrances and internal dimensions. Discriminant function analysis produced parsimonious models for each site, with high rates of correct classification, based on structural, floristic and breeding success parameters. Lack of transference of predictive powers of the models between sites demonstrates the apparent variability in hornbill nesting behaviour and indicates the need for caution when extrapolating models to other sites. Possible factors affecting the nesting density and distribution of each hornbill species are discussed.     

Nest site selection by yellow-faced honeyeaters Lichenostomus chrysops

When choosing subsequent nest sites, species that produce multiple broods in a single breeding season have the option of dispersing from a site where they previously suffered depredation, i.e., a predator-avoidance tactic. In our study on yellow-faced honeyeaters Lichenostomus chrysops it was found that 89% of nest failures were attributed to nest depredation, the primary cause of reproductive failure. Pairs re-nested further from depredated nesting attempts than from successful nesting attempts and progressively higher above the ground as the breeding season progressed. Pairs nesting in dioecious Coprosma quadrifida plants only nested in non-fruiting male plants. Artificial nests were used to test the hypothesis that nest height and plant preferences were strategies to reduce the risk of depredation. There was no evidence that either higher nests or nests in non-fruiting C. quadrifida achieved reduced levels of depredation during 14 days of artificial nest exposure. Specific nest site characteristics were not found to be associated with nest outcome for either natural or artificial nests. Our study provides further evidence that species may choose a diverse range of nest sites in order to avoid predators from developing specific search images and then, following depredation, compensate by rapidly re-nesting away from the failed attempt.     

Effect of soil hardness on aggression in the solitary wasp Mellinus arvensis

1. Two alternative nesting strategies are exhibited by soil‐nesting Mellinus arvensis females, digging a new nest (diggers) and searching for an old unoccupied burrow (searchers). Wasps appear unable to distinguish between occupied and unoccupied nests, and aggressive interactions between searchers and nest owners at nest entrances are frequent. 2. In aggressive encounters, there is an advantage in size and residency status. 3. The costs associated with the two nesting strategies vary across geographically separated populations: nest digging incurs costs in terms of time, and these vary according to the hardness of the soil substrate; nest searching is variably costly in terms of risk of injury in aggressive encounters with nest‐owning females. 4. Individual female wasps can switch between nesting strategies, and thus soil hardness, by affecting the cost of nest construction, affects the relative frequencies of the two nesting strategies within a population, favouring an increase in the searching strategy. This, in turn, affects the frequency and intensity of aggression between females at a nesting aggregation. 5. Female body size is correlated with soil hardness. As aggressive encounters are more frequent in sites with hard soil substrates, there is increased selective advantage in having large body size at these sites. 6. Body size is determined primarily by the availability of food resources during larval development, which is, to a degree, a function of the size of the adult female. There is a trade‐off between provisioning a few cells with many provisions in each, leading to the development of few but large adults, as opposed to many cells with few provisions, leading to many small offspring. The relative advantage of these two provisioning strategies is, at least in part, a function of the hardness of the soil substrate.     

Philopatry, nest choice, and aggregation temporal–spatial change in the digger wasp Cerceris arenaria (Hymenoptera: Crabronidae)

Studying temporal and spatial changes of aggregations of digger wasps through nesting seasons is interesting because of its link to social evolution via a semi-social pathway, but information about this topic is scarce. An aggregation of Cerceris arenaria Latreille was studied during 1997–1999 and for a short period in 2001. The colony grew in area and number of nests through the first three seasons, but the aggregation reduced in 2001, and this decrease was confirmed by a count of nests made in 2003 and 2005. The positions of the nest entrances remained almost constant over the years, at least in high-nest-density areas, because of the repeated use of the same burrows from which they emerged. This fact permits the population to exploit for many years a restricted area and could act as a nest-density-regulation method for fossorial species that rarely dig new nests, preventing an excessive nest density. The newly emerged females did not dig new nests, showed a high philopatry in the choice of their first nest, and preferred to look for other nests to occupy close to the ones recently abandoned (mainly for an undergone usurpation by a conspecific female). Philopatry, along with lack of new nest digging and position-dependent nest choice, is probably an important factor maintaining spatial stability of the aggregation through the years and could represent a first step in the evolution of a higher sociality in apoid burrowing Hymenoptera.     

ADAPTATIONS TO NESTING HABITAT IN THE REPRODUCTIVE BEHAVIOUR OF THE BLACK-BILLED GULL LARUS BULLERI

SUMMARY The nesting habitats adopted by most colonies of Black-billed Gulls Larus bulleri are river-beds that are subject to flooding. A number of respects in which the reproductive behaviour of Black-billed Gulls differs from that of at least most other gulls, such as Black-headed Gulls, can be viewed as adaptations, or byproducts of adaptations, to such nesting habitats:—

• (a) A different breeding site from the year before is often selected
• (b) The bulk of pair formation is accomplished before the gulls occupy their breeding sites; nesting territories are set up by mated pairs
• (c) In hostile encounters during the pair formation phase, site attachments are weak or transitory
• (d) In agonistic situations generally, attack thresholds appear to be relatively high, and fleeing thresholds relatively low
• (e) High intensity forms of“Choking” appear to be missing from the agonistic display repertoire
• (f) The growth of nest groups is rapid and orderly; nesting territories are small and the concentration of nests in the groups high
• (g) The time between occupation of the gullery site and the start of laying is short, and the synchrony of laying is high; this is probably related to the close proximity of the nests
• (h) The parents and young abandon the nest very soon after the egg hatch, and no “brood” nests are made
• (i) The young may develop locomotory powers more quickly than is the case in most other species; they can swim at an early age, and take to the water in tightly packed groups during alarms

The question of why these gulls should choose such vulnerable breeding sites is discussed.     

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.     

Raptors breeding on weaver nests in trees and on man-made structures

Several raptor species nest on top of large weaver nests. These weaver nests are usually sited in trees, but 11.7% of Red-billed Buffalo-Weaver Bubalornis niger and 25.7% of Sociable Weaver Philetairus socius nests occur on man-made structures. In an extensive literature search, a total of 16 raptor species were recorded as nesting on top of weaver nests. At least 10 raptor species used weaver nests built in trees. Seven raptor species used weaver nests on man-made sites and four raptor species only used weaver nests built on man-made sites. No owls have been recorded as using weaver nests on man-made sites. There are historical records of raptors nesting on top of weaver nests in trees, while nesting on top of weaver nests sited on man-made structures appears to be a more recent adaptation. Costs and benefits of nesting on man-made sites are briefly listed. Nesting on man-made sites may increase (by both weavers and raptors) and raptor researchers are encouraged to document cases of raptors nesting on weaver nests where these are placed in trees or artificial sites, so that there is a record of changes in frequency of nest site usage by raptors.