Intraspecific nest parasitism in the sand waspStictia heros (Fabr.) (Hymenoptera: Sphecidae)

Two forms of intraspecific nest parasitism are described for the neotropical sand waspStictia heros (Fabr.). Females behaving aggressively made aerial attacks on prey-laden females as they approached their nest for provisioning. Attacks resulted in prolonged grappling on the sand, and the relative sizes of the contestants influenced final possession of the prey. Also, the probability that returning females were attacked was directly related to the size of the prey being delivered. Females displaying marauding behavior entered the nests of other females, removed prey, and deposited it in their own nest. Females usually raided nests that were near their own nest and tended to concentrate raiding attempts on nests that previously yielded prey. We also describe the behavior associated with the delivery of hunted prey and examine possible interrelationships between wasp size, prey size and hunting trip duration.     

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.     

Parasite presence acts as a proximate cue in the nest-site selection process of the solitary digger wasp,Ammophila dysmica (Hymenoptera: Sphecidae)

Nest-site selection by the solitary ground-nesting wasp Ammophila dysmicaMenke was studied during 1982–1986 in California's Sierra Nevada Mountains. Digging females detected and discriminated among members of a complex of insect intruders, including a number of predators and parasites. In response to some natural enemies, wasps sometimes permanently abandoned nests under construction. These selective nest abandonments reduced the mean number of cleptoparasitic chrysidid wasps, Argochrysis armillaBohart, attending the excavation of completed nests. The number of A. armillaattending a nest excavation correlated positively with the probability of nest parasitism by this species. Selective nest abandonment may reduce parasitism rates.     

Sex-differences in nest defence in Fieldfares Turdus pilaris in relation to their size and physical condition

The nest defence of solitary-breeding Fieldfares Turdus pilaris against a human intruder, a corvid and a hawk was studied in a mountain birch forest in central Norway in 1990–2002. Females defended their nests against a human intruder more vigorously than did males before hatching, and vice versa after hatching. Although both sexes increased their nest defence, only males showed a significant increase and showed a greater defence than females at the chick stage. Females with 1-week-old nestlings had longer wings, were heavier and in a better physical condition than those that failed to get nestlings to that stage. Females, but not males, lost condition during nesting. Birds in better condition were more aggressive. The pattern of nest defence by males and females against the corvid was similar to that shown against human intruders, whereas males took greater risks than females towards the raptor. There was weak evidence that birds that were more successful showed greater levels of nest defence. The sex-role reversal in nest defence may be connected to differences in costs in relation to the renesting potential and to the physical condition of the sexes.     

Does Conspecific Attraction Affect Nest Choice in Budgerigars (Melopsittacus undulatus: Psittacidae: Aves)?

In their native Australia the budgerigar's breeding season is very short and rapid nest establishment is crucial. To minimize the time required to locate and establish nesting sites, female budgerigars were predicted to use cues on nest site locations provided by other females in their colony. Experimental results from captive birds indicated that, while females can be attracted to nests that are occupied by other females, the social and breeding experience of individuals affects this use of conspecific cues. Females that have had altercations with other females that resulted in physical injuries showed an aversion to apparently occupied nestboxes. Females that had no such social experience prior to testing were attracted to apparently occupied nestboxes. Breeding experience, as well as social experience, may have an effect on the use of conspecifics as cues. Experienced breeding females were less likely to be attracted to apparently occupied nestboxes than were females that had never bred before.     

Comparative study of the nesting behaviour of Tachysphex inconspicuus (Kirby) (Hymenoptera: Crabronidae) in two locations in southeast Brazil

The nesting behaviour of the solitary wasp Tachysphex inconspicuus (Kirby) was comparatively studied based on observations made in two environmentally distinct locations in southeast Brazil: upper portions of two sandy beaches and a dirt road in a forested area. Motor patterns related to nest construction were similar in all observed wasps, but some behavioural features, which were different among the observed wasps in the two environments, seem to be constrained by ecological factors. T. inconspicuus can be behaviourally differentiated from several other species of the genus by the following combination of features: they close the nest temporarily during its provision, do not let the excavated sand to form into a mound in front of the nest entrance, construct unicellular nests, and provision the nest with cockroaches. Moreover, females T. inscospicuus use their hind legs in a peculiar way to manipulate the prey, which allows them to associate the use of the mandibles in prey transport with the possibility of excavating the temporary plug and entering the nest without releasing the prey. Chorisoneura lopesi Albuquerque, C. excelsa Albuquerque & Silva, and unidentified species of Riatia (Blattodea: Blatellidae) were found as prey and Amobia floridensis (Towsend) (Diptera: Sarcophagidae) as parasitoid of T. insconspicuus.     

Kin grouping is insufficient to explain the inclusive fitness gains of conspecific brood parasitism in the common eider

Conspecific brood parasitism allows females to exploit other females' nests and enhance their reproductive output. Here, we test a recent theoretical model of how host females gain inclusive fitness from brood parasitism. High levels of relatedness between host and parasitizer can be maintained either by: (a) kin recognizing and parasitizing each other as a form of cooperative breeding or (b) natal philopatry and nest site fidelity facilitating the formation of kin groups, thereby increasing the probability of parasitism between relatives nesting in close proximity. To address these two hypotheses we genotyped feathers and hatch membranes of common eiders (Somateria mollissima) from western Hudson Bay, Canada, using a noninvasive sampling methodology. We found that most instances of brood parasitism do result in inclusive fitness gains. Furthermore, females with failed nests moved an average of 492 m from their previous year's nest site, while successful females only moved an average of 13 m. Therefore, we observed host–parasite relatedness can occur at levels higher than would be expected by chance even in the absence of kin grouping, suggesting that closely related females nesting near one another is not essential to maintain high host–parasitizer relatedness. In addition, kin grouping is only a transient phenomenon that cannot occur every year due to the propensity for females of failed nests to nest farther away from their nest site in subsequent years than females with successful nests, which provides support for kin recognition as a more likely mechanism to maintain high host–parasitizer relatedness over time.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

Anti-social and social behaviour of adolescent yellow-rumped caciques (Icterinae: Cacicus cela)

In an individually marked population in southeastern Peru, males and females of the polygynous, colonial yellow-rumped cacique delayed successful breeding until they were 3 years old. Adolescent (2-year-old) females competed for nest sites, built nests, laid fertile eggs, incubated, and sometimes fed nestlings, but always abandoned nests before they fledged young. Females that completed nests during their adolescent years were also more successful in establishing nest sites the following year. Adolescent females that failed to establish nest sites and those that lost nests to predators tended to leave the study area altogether. By attempting to nest early, adolescent females may facilitate future nesting attempts by forming coalitions with other females and assessing nest site quality in an area. Adolescent males, on the other hand, avoided aggressive interactions with other males, but were very aggressive towards adult females and fledglings. Adolescent males attacked females in nests, chased females to and from the colony, pecked and mounted recent fledglings, and, rarely, pulled nestlings out of nests. This harassment resulted in the death of nestlings and fledglings and was energetically costly to females, which often fought adolescent males while defending their young. Adolescent males preferentially harassed nestling-feeding females, avoided egg-laying females and were never observed attempting to mate with females. Adult males rarely interfered during any of these interactions. Such anti-social behaviour may function as a way of practising adult behaviour, as a means of eliminating future competitors, or may be an epiphenomenon of rising hormone levels and therefore of no adaptive significance.     

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

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

Molecular identification of brood‐parasitic females reveals an opportunistic reproductive tactic in ruddy ducks

In many taxa, females lay eggs in the nests of other conspecifics. To determine the conditions under which conspecific brood parasitism develops, it is necessary to identify parasitic offspring and the females who produce them; however, for most systems parasitism can be difficult to observe and most genetic approaches have relatively low resolving power. In this study, we used protein fingerprinting from egg albumen and 10 microsatellite loci to genetically match parasitic ducklings to their mothers in a population of ruddy ducks (Oxyura jamaicensis). We found that 67% of nests contained parasitic offspring, and we successfully identified their mothers in 61% of the cases. Of the parasitic females identified, 77% also had nests of their own (i.e. a dual tactic, where females both nest and lay parasitically), and we found no evidence that parasitic females pursued a specialist (parasitism only) tactic. We also found that parasitic egg laying was not influenced by nest loss, predation or female condition. Thus, in contrast to most waterfowl studied to date, female ruddy ducks appear to lay parasitic eggs whenever the opportunity arises.     

Common Cuckoos Cuculus canorus change their nest‐searching strategy according to the number of available host nests

In recent decades, numerous studies have examined factors affecting risk of host nest parasitism in well‐known avian host–parasite systems; however, little attention has been paid to the role of host nest availability. In accordance with other studies, we found that nest visibility, reed density and timing of breeding predicted brood parasitism of Great Reed Warblers Acrocephalus arundinaceus by the Common Cuckoo Cuculus canorus. More interestingly, hosts had a greater chance of escaping brood parasitism if nesting was synchronized. Cuckoo nest searching was governed primarily by nest visibility at high host‐nest density. However, even well‐concealed nests were likely to be parasitized during periods when just a few hosts were laying eggs, suggesting that Cuckoos adjust their nest‐searching strategy in relation to the availability of host nests. Our results demonstrate that host vulnerability to brood parasitism varies temporally and that Cuckoo females are able to optimize their nest‐searching strategy. Moreover, our study indicated that Cuckoos always manage to find at least some nests to parasitize. Thus, in this case, the co‐evolutionary arms race should take place mainly in the form of parasitic egg rejection rather than via frontline pre‐parasitism defence.     

Aphid-hunting wasps: a field study of Passaloecus

• 1 Three species of pemphredonine wasps, Passaloecus insignis, P.gracilis and P.corniger, were observed nesting in disused beetle holes in pine boards. The species differed from one another in the size, height above ground and aspect of the holes selected as nest sites.
• 2 P.insignis and P.gracilis stocked their nests with aphids of several species, but P.corniger females stole paralysed aphids from nests stocked by P.insignis, P.gracilis or by conspecifics, sometimes breaking open the outer door of a nest to do so.
• 3 The outer doors of the nests were of pine resin; P.corniger and P.insignis added pellets of other material to the surface. P.corniger females revisited their nests after sealing the outer door and behaved as if scent-marking them, perhaps conferring protection against robbery by conspecifics.
• 4 Adult male wasps probably fed on honeydew. They appeared early and seemed to have a flying season of only a few days, whereas females probably lived and flew for about eight weeks after emergence.
• 5 The behaviour of nest-stocking female wasps is considered in relation to the temperature of their microhabitats.
• 6 The possible use of pemphredonine wasps for the biological control of aphids or thrips in glasshouses is discussed.

     

Decoding dumping ducks

Conspecific brood parasitism, where females of the same species lay eggs in each other's nests, is common in waterfowl, and is usually considered costly to host females, which are stuck looking after eggs and chicks that are not their own. However, since female waterfowl often exhibit an unusual propensity to nest near where they were born, there has been some uncertainty over whether, in ducks and geese, laying in nests of conspecifics really is parasitism. Do parasitic and host females tend to be related? And is parasitism actually a form of cooperation in disguise? In a population in Hudson Bay, Andersson & Waldeck (this issue) found that ‘parasitic’ eggs in nests of the common eider, Somateria mollissima sedentaria, are more closely related to host eggs than expected by chance. In fact, host and ‘donor’ eggs are more closely related than are females breeding at neighbouring nests. The Hudson Bay population of common eiders is unusual, because unlike in more benign climates, females do not tend to breed near their natal nest. Spatial proximity alone cannot account for the high relatedness between host eggs and ‘dumped’ or donor eggs. Instead, the high relatedness values are probably the result of active recognition, where females favour kin, either when dumping or accepting eggs. These new data, along with evidence indicating that the donor lays the first egg in the nest nearly half the time, suggest that what appears to be parasitism in common eiders may be a form of kin‐based cooperation.     

Sex-specific defence behaviour against brood parasitism in a host with female-only incubation

Nest protection against intruders is an indispensable component of avian parental care. In species with biparental care, both mates should evolve nest defence behaviour to increase their reproductive success. In most host-parasite systems, host females are predicted to have more important roles in nest defence against brood parasites, because they typically are primarily responsible for clutch incubation. Male antiparasitic behaviour, on the other hand, is often underestimated or even not considered at all. Here we investigated sex-specific roles in four aspects of great reed warbler (Acrocephalus arundinaceus) nest defence against a brood parasite—the cuckoo (Cuculus canorus), namely (1) mobbing, (2) nest attendance/guarding, (3) nest checking and (4) egg ejection. Using dummy experiments, simulating brood parasitism and by video-monitoring of host nests we found that males took the key roles in cuckoo mobbing and nest guarding, while females were responsible for nest checking and egg ejection behaviours. Such partitioning of parental roles may provide a comprehensive clutch protection against brood parasitism.     

An alternative strategy for maintenance of eusociality after nest destruction: new nest construction in a primitively eusocial wasp

Although nests are central to colonial life in social insects, nests are sometimes damaged by predators or natural disasters. After nest destruction, individuals usually construct new nests. In this case, a sophisticated mechanism like the scent trail pheromone used in large insect colonies that recruit individuals to new nest sites would be important for the maintenance of eusociality. In independent-founding Polistes wasps, it is well known that queens enforce workers physiologically on the natal nests even if evidence of trail pheromone use has not been exhibited. We investigated the effect of the queen on an alternative strategy for the maintenance of eusociality by first females after nest destruction in the primitively eusocial wasp Polistes chinensis. We predicted that the first females in queen-absent colonies have various behavioral options after nest destruction. Even if the females construct new nests cooperatively with other individuals, the new nest construction should be conducted more smoothly in queen-present colonies because the queens regulate the behavior of wasps. We made wasps construct new nests by removing the entire brood from existing nests. The presence of the queen did not cause variation in the alternative strategy of the first females, as the first females (workers) usually constructed new nests cooperatively irrespective of the queen-presence. Thus, the workers in the queenpresent colonies affiliated to the new nest construction more smoothly and constructed new nests more efficiently than workers in the queen-absent colonies. Our results suggest that the presence of the queen is important for maintaining eusociality in primitively eusocial wasps after nest destruction. Received 8 February 2005; revised 5 October 2005; accepted 17 October 2005.     

Interactions between the Social Digger Wasp, <Emphasis Type="Italic">Cerceris rubida</Emphasis>, and Its Brood Parasitic Flies at a Mediterranean Nest Aggregation

The fitness consequence of many behaviours of the small digger wasp, Cerceris rubida (Hymenoptera: Crabronidae), the only European species of its genus in which females share nests, are still unknown. Here, I present novel data on the nesting patterns and nest parasites of an Italian population of this wasp, with emphasis on which behavioural strategies may have evolved to reduce brood parasitism. Nests were established mainly in horizontal surfaces with scarce vegetation and hard soil, resulting in spatially clumped nests; the extent of nest aggregation increased over a 6-year period. Wasp brood cells are attacked by the miltogrammine fly Pterella grisea (Diptera: Sarcophagidae), which waits for nest-returning wasps on perching sites and then follows them in flight (satellite flight), eventually landing on the nest entrance and larvipositing without entering further in the tunnel. This technique seems to be adaptive for the parasitic flies, which would be rejected from nests by the guarding wasps if attempting to enter. The daily activity of the flies closely matched the host wasp’s provisioning activity, but C. rubida females were able to partially confound the tracking flies by performing evasive manoeuvres while returning to the nest. Patches with higher nest density and nests with more resident females did suffer more fly landings on nest entrances (a prerequisite for larviposition). These trends, however, disappeared on a per nest basis and on a per wasp provisioning flight basis, respectively. Across two years, only 6% of brood cells were parasitized, a picture similar to what observed for miltogrammine flies at nest aggregations of other Cerceris spp., and new data are necessary to test if there is a benefit of either nest density or nest sharing against P. grisea parasitism.     

Rediscovery of the workerless inquiline antPogonomyrmex colei and additional notes on natural history (Hymenoptera: Formicidae)

Summary Pogonomyrmex colei is a workerless inquiline ant known only from nests ofP. rugosus, its closest relative. Ten of 776 (1.3%) host nests were parasitized at a site in central Arizona, while none of 1499 potential host colonies were parasitized at two other locales. Colonies ofP. colei are perennial, and host alate females in 9 of 10 colonies demonstrates that host queens survive parasitism. Three of 10 colonies died over 19 colony years of observation, while only 1 of 601 colonies became newly parasitized. Mating occurs in morning for up to 2–3 days following summer and fall rains and in afternoon during cool fall days. Mating is intranidal just outside the nest entrance, with males returning to the natal nest. MaleP. colei may be flightless because their wing area is reduced compared to host males. Females fly from the nest and locate potential host colonies by following trunk trails. Workers are the largest barrier to nest establishment, as they removed over 90% ofP. colei females placed in trunk trails or that entered host nests. Males and females ofP. colei andP. anergismus, the only other congeneric inquiline species, are diminutive compared to their hosts, with females 30% lighter than host workers. Fat content is lower and water content is higher inP. colei andP. anergismus females than in their hosts.     

Factors affecting the acceptance of alien conspecifics on nests of the primitively eusocial wasp,Ropalidia marginata (Hymenoptera: Vespidae)

In the primitively eusocial wasp, Ropalidia marginata,individual females are known to drift from one newly founded nest to another. In the laboratory, young (<6- to 8-day-old) alien wasps are accepted onto unrelated colonies, while older (>6- to 8-day-old) wasps are not. Here we have investigated the factors that could influence the acceptance of foreign conspecifics onto unrelated nests. Individually marked wasps of different ages, isolated immediately after eclosion from the natal nest and from each other, were introduced onto unrelated recipient nests. Considered separately, both age and ovarian condition seemed to influence the probability of acceptance as well as the levels of aggression and tolerance received by the introduced wasps. However, partial correlation analysis and multiple regression analysis indicated that only age had a direct influence and that the ovarian condition acts only through age, a variable with which it is highly correlated. The observed acceptance of young aliens and rejection of old aliens are less likely to be due to the perception of older wasps as a reproductive threat rather than some age-related factor, other than ovarian condition, for example, the relative ease with which younger wasps can be molded into desired roles.     

Brent goose <Emphasis Type="Italic">Branta bernicla bernicla</Emphasis> feeding behaviour during incubation,Taïmyr Peninsula,Russia

Incubating birds must balance the time and the energy invested in incubation with the energy acquisition for their survival. Many factors such as weather and predation influence this trade-off. In Arctic geese, only females incubate, and they leave the nest regularly to feed while males invest in keeping their nests and mates safe. This study conducted on Big Bird Island (Taïmyr Peninsula) during the summer of 2004 examined the incubation behavior of dark-bellied brent geese Branta bernicla bernicla to assess the effect of date, period of day and weather conditions on the incubation and feeding behaviors of females and males. Females were at their nests only for 65% of the total time observed. This very low value, compared to other goose species, could be the result of the combined effects of good weather conditions, low predation pressure and opportunities to feed close to the nest. We found differential adjustments of male and female behaviors. Females appeared to focus on the trade-off between feeding and incubating, in relation to weather conditions, and on their own energy balance. Males appeared to respond primarily by the absence of the female from their nest.