Behaviorally mediated spatial and temporal refuges from a cleptoparasite,Argochrysis armilla (Hymenoptera: Chrysididae), attacking a ground-nesting wasp,Ammophila dysmica (Hymenoptera: Sphecidae)

Summary It has been suggested that parasite pressure favors the evolution of sociality within the Hymenoptera. I analyzed the impact of a chrysidid nest parasite, Argochrysis armilla, on its host, the solitary ground-nesting wasp, Ammophila dysmica, to assess the role of parasitism in favoring two steps towards sociality: aggregated nesting and delayed nest provisioning. The foraging strategy of Argochrysis armilla involves discovering host nests during excavation, learning the locations of discovered nest, and returning to oviposit in nests during nest provisioning; I therefore assessed the influence of host behavior on (1) parasite discovery of nests and (2) parasite oviposition in nests. Significantly fewer parasites discovered host nests that were excavated during the early morning and late afternoon, due to partial asynchrony of host and parasite activity. Nests excavated in areas of low nest density were also less likely to be discovered; use of low density nest sites increased during periods of high parasite activity. Due to a rapidly decelerating rate of parasite recruitment to nests under excavation, the duration of nest digging had only a limited influence on nest discovery by parasites. The probability of parasite oviposition in a host nest was determined by the number of parasites discovering the nest during excavation and by the time between nest excavation and provisioning; delaying nest provisioning reduced the risk of parasite oviposition. Delayed provisionings primarily appeared to be a result of the stochastic process of hunting and prey encounter. The number of provisions placed in a nest (one vs two) had no effect on the probability of nest parasitism. Spatial patterns of parasitism were directly density dependent in 1984 and density independent in 1986. In this system parasite pressure acts against the formation of nesting aggregations and in favor of delayed nest provisioning. The dependence of these results on species-specific aspects of the parasite's foraging strategy and the host's defensive strategy suggests, however, that different parasite species may generate qualitatively different selection pressures, potentially contributing to the diversity of nesting behavior in the Hymenoptera.     

Patterns of Predation Risk and Survival of Bird Nests in a Chilean Agricultural Landscape

Abstract: We used experimental nests baited with California Quail (  Callipepla californica ) eggs or clay eggs to examine relative risks of nest predation in an agricultural landscape and in two large forest preserves in a south-temperate rainforest in Chile. The most common predators, as identified by marks on clay eggs, were a caracara (   Milvago chimango ), a blackbird ( Curaeus curaeus ), and rodents. Nest losses from predation were similar in large and small forest patches and lower in patches than in extensive forest. In general, predation risk was higher (and nest survival therefore lower) on forest edges than in forest interior, in short-grass pasture than in tall-grass pasture, in narrow corridors than in wide corridors, and on visible nests than on concealed nests. High predation risks in pasture habitat tended to increase the risk of nest predation in adjacent forest edges. For open-cup nesters, the risk of nest predation was relatively high in the present agricultural landscape, indicating that much of the available wooded habitat (  forest edges, narrow corridors) offers poor nesting habitat, although it may be suitable for foraging and traveling. The numerous bird-plant mutualisms in this landscape may be at risk if nesting success of the principal mutualists is consistently low.     

The function of polydomy: the ant <Emphasis Type="Italic">Crematogaster torosa</Emphasis> preferentially forms new nests near food sources and fortifies outstations

Many ant species are polydomous, forming multiple spatially segregated nests that exchange workers and brood. However, why polydomy occurs is still uncertain. We investigated whether colonies of Crematogaster torosa form new polydomous nests to better exploit temporally stable food resources. Specifically, we tested the effect of food presence or absence and distance on the likelihood that colonies would form a new nest. Because this species also forms little-known structures that house only workers without brood (outstations), we also compared the function of this structure with true nests. Laboratory-reared colonies were connected to a new foraging arena containing potential nest sites with or without food for 4 months. When food was present, most colonies formed polydomous nests nearby and the remainder formed outstations. When food was absent, the behavior of colonies differed significantly, frequently forming outstations but never polydomous nests. Distance had no effect on the type of structure formed, but when food was present, a larger proportion of the workforce moved shorter distances. Workers often fortified the entrances to both structures and used them for storage of dried insect tissue (“jerky”). In an investigation of spatial fidelity, we found that workers on the between-nest trail were associated with the original nest, whereas workers collecting food were more likely to be associated with the new nest or outstation. C. torosa appears to have a flexible colony structure, forming both outstations and polydomous nests. Polydomous nests in this species were associated with foraging and were only formed near food resources.     

Nest concealment and parental behaviour interact in affecting nest survival in the blackcap (Sylvia atricapilla): an experimental evaluation of the parental compensation hypothesis

Nest concealment varies strongly within populations of many species. Although some studies have revealed the beneficial effects of concealment in mitigating predation pressure on nests, other studies were unable to find similar effects. One potential reason for the mixed results is that parental behaviour may compensate for the effects of nest cover, and specifically designed experimental studies are needed to reveal this compensation. I studied the effects of concealment on the probability of nest survival in the blackcap (Sylvia atricapilla), by experimentally manipulating the degree of nest-foliage cover. There was a significant effect of the treatment depending on nest type and the phase of nesting. Whereas there was no effect of concealment on nest survival in natural nests, there was a positive effect in real nests baited with plasticine clutches (i.e. without parental activity). Parents probably behaviourally compensated for poor concealment in natural nests (nest guarding, defence). In line with this, there was no effect of concealment on nest survival during incubation, whereas there was probably a positive effect in the nestling phase. Parents spent more time on the nest during incubation (80%) than during the care of nestlings (40%) and, consequently, had more opportunities to compensate for poor cover. In general, we cannot use single measures of behaviours or states (nest concealment) as an indication of predation risk because of the capacity for compensation in other behaviours.Communicated by C. Brown     

Potential Impact of Mammalian Nest Predators on Endemic Forest Birds of Western Mauna Kea, Hawaii

I used dummy nest experiments to investigate the role of nest predation by introduced mammals asa significant limiting factor for the endangered Palila ( Loxioides bailleui ) and other endemic birds on the western slope of Mauna Kea. Overall predation rates on dummy nests were extremely low. Rates were comparable to those on actual Palila nests, indicating that dummy nests give a valid representation of the dynamics of nest predation. The black rat ( Rattus rattus ) was the only important predator. Feral cats ( Felis catus ) played only a minor role, and there was no evidence to implicate house mice ( Mus musculus ) as nest predators. Four factors appear to be responsible for the low nest predation rates: (1) only a single species of predator (black rat) is involved; (2) rat densities are extremely low on Mauna Kea; (3) low prey (nest)densities preclude density-dependent predation, and (4) rats have alternative foods that are more abundant and accessible than arboreal bird nests. Since mammalian predation appears unlikely to have a significant impact on the Palila, other factors must be limiting the abundance of this endangered species.     

Patterns of Nest Predation on Artificial and Natural Nests in Forests

Abstract:  Artificial nest experiments have been used in an attempt to understand patterns of predation affecting natural nests. A growing body of literature suggests that neither relative rates nor patterns of predation are the same for artificial and natural nests. We studied nest predation and daily mortality rates and patterns at real and artificial ground and shrub nests to test the validity of artificial nest experiments. We monitored 1667 artificial and 344 natural nests, over seven trials, in three regions, across 58 sites in Ontario. We controlled for many of the factors thought to be responsible for previously reported differences between predation rates on natural and artificial nests. Although artificial nests in our study resembled natural nests, contained eggs of appropriate size, shape, and color of target bird species, and were placed in similar microhabitats as natural nests, the rates of predation on these nests did not parallel rates on natural nests for any region in terms of absolute rate or pattern. Predation rates on artificial nests did not vary between years, as they tended to for natural nests, and the magnitude of predation pressure on artificial ground nests compared with shrub nests did not show the same pattern as that on natural nests. In general, rates of predation on artificial nests were significantly higher than on natural nests. Our results suggest that conclusions derived from artificial nest studies may be unfounded. Given that many influential ideas in predation theory are based on results of artificial nest experiments, it may be time to redo these experiments with natural nests.     

Parasitic common goldeneye (<Emphasis Type="Italic">Bucephala clangula</Emphasis>) females lay preferentially in safe neighbourhoods

Nest predation has been suggested as an explanation of the adaptive significance and evolution of conspecific brood parasitism, an alternative reproductive tactic pursued by females in several animal taxa. I used new nest boxes that contained only decoy eggs and were erected on lakes differing in real nest predation risk to test this hypothesis in the common goldeneye (Bucephala clangula), a hole-nesting duck. I used broken eggs to simulate predation risk of the boxes to determine if parasites having no previous experience with the boxes discriminate between seemingly safe and risky nest sites. Parasites laid eggs in the experimental boxes independently of the simulated predation risk, suggesting that they do not use broken eggs or nest disarray as indicators of predation intensity. Parasites preferred experimental boxes on lakes where real nest predation risk was low, supporting the nest predation risk hypothesis. Assuming that females in high risk areas have had experience of nest predation, they may take this into account in selecting host nests.     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.     

The evolution of worker–queen polymorphism in Cataglyphis ants: interplay between individual- and colony-level selections

In many ants, young queens disperse by flying away from their natal nest and found new colonies alone (independent colony founding, ICF). Alternatively, in some species, ICF was replaced by colony fission, in which young queens accompanied by workers found a new colony at walking distance from the mother nest. We compared the queen morphology of Cataglyphis floricola, which disperses by fission, with that of its most likely living ancestor, Cataglyphis emmae, which disperses by ICF. As in other species, the transition from ICF to fission is associated with queen miniaturization. Interestingly, C. floricola presents two types of small queens: brachypters (with short non-functional wings) and ergatoids (worker-like apterous queens). Ergatoids are, on average, 2.8 mg lighter and have half the number of ovarioles than brachypters, which limits the advantage for a colony to produce ergatoids instead of brachypters. Furthermore, more ergatoids are produced than brachypters, but their individual survival rate is lower. During colony fission, 96% of the cocoons containing brachypters but only 31% of those containing ergatoids are transferred to the daughter nests where, after emergence, they compete for becoming the next queen. The remaining queen cocoons, which stay in the mother queen's nest, are eliminated by workers upon emergence, probably to maintain monogyny. This waste of energy suggests that producing ergatoids instead of brachypters is unlikely to increase colony efficiency. We argue that the evolution of ergatoids could derive from a selfish larval strategy, developing into worker-like queens in spite of the colony interest.     

Dispersal decisions and predispersal behavior in <Emphasis Type="Italic">Polistes</Emphasis> paper wasp ‘workers’

Social insects are popular models for studying the evolution of cooperation. Casteless taxa where individuals have the flexibility to either nest alone or cooperate are particularly valuable for understanding the causes and consequences of cooperative behavior. For example, some ‘workers’ from Polistes paper wasp nests disappear from their natal colony soon after pupal emergence and nest independently. However, little is known about dispersal behavior. In this paper, I compare predispersal behavior of wasps who leave their natal colony soon after emergence with behavior of individuals who remain on the natal colony as true workers. I found that P. dominulus females with short nest tenure behave much like gynes (reproductive-destined offspring produced at the end of the season), as wasps with short nest tenure are behaviorally selfish while on the natal colony. They spend a smaller proportion of their time foraging and a larger proportion of their time resting than workers with long nest tenure. In addition, I assessed the factors that may favor early dispersal. Nest environment strongly influenced dispersal; large colonies had a smaller proportion of females with short nest tenure. Queen turnover also increased dispersal behavior perhaps because queen turnover reduces relatedness between a colony’s current and future offspring, thereby reducing the kin-selected benefits of cooperation. Therefore, casteless social insects exhibit a surprising degree of reproductive flexibility. Individuals may use information about their internal state and nest environment to optimize their behavioral strategies.     

The effect of incubation temperature on hatchling quality in the olive ridley turtle, Lepidochelys olivacea, from Alas Purwo National Park, East Java, Indonesia: implications for hatchery management

Nest protection through egg relocation from natural nests into protected hatcheries is a common practice used at rookeries around the world to increase hatchling recruitment into sea turtle populations. However, rarely have the impacts of this practice on hatchling recruitment and quality been assessed. This study investigated the influences of the thermal nest environment of olive ridley turtles Lepidochelys olivacea on emergence success and quality of hatchlings of hatchery nests in Alas Purwo National Park, East Java, Indonesia (2009 and 2010 nesting seasons). Nest temperatures above 34?°C for at least 3 consecutive days during incubation in the hatchery resulted in decreases in emergence success and locomotor performance of hatchlings. The use of the hatchery is recommended due to extremely high predation rate of nests left on the beach; however, altering hatchery management practice by spacing nests one meter apart and providing shade should improve hatchery outcomes now and into the future.     

Chemical mimicry in the root aphid parasitoid Paralipsis eikoae Yasumatsu (Hymenoptera: Aphidiidae) of the aphid-attending ant Lasius sakagamii Yamauchi & Hayashida (Hymenoptera: Formicidae)

Summary. An aphidiid wasp, Paralipsis eikoae, was associated with both Lasius niger and L. sakagamii attending the wormwood root aphid Sappaphis piri. An L. sakagamii worker was observed carrying a winged female P. eikoae to its nest with its mandible, but it did not kill the wasp. Once accepted by the ants, the wasp often mounted and rubbed against the worker ants and sometimes teased them to regurgitate food to itself. No workers in the colony attacked the wasp. Conspecific foreign workers, however, viciously attacked the wasp when encountered. Gas chromatography-mass spectrometry analyses showed that the accepted wasp had complex cuticular hydrocarbons that were very similar to those of its host ants, whereas the winged wasps collected outside the ant nest showed only a series of n-alkanes. Additionally, the accepted wasp had a hydrocarbon profile closer to that of its host ants than to the conspecific foreign ants. We believe the wasp mimics ant cuticular hydrocarbons to integrate into the ant nest, acquiring the hydrocarbons by mounting and rubbing against the ants. In contrast, the cuticular hydrocarbons of the emerged wasp contained larval and pupal hydrocarbons of L. sakagamii that were also similar to those of L. niger. Both ant species rejected adult workers of the other species but accepted their larvae and pupae. We suggest that the emerged P. eikoae mimics the cuticular hydrocarbons of these Lasius larvae and pupae, which allows P. eikoae to be accepted by both L. sakagamii and L. niger. Received 11 March 1998; accepted 22 July 1998.     

Nestmate recognition in three species of stenogastrine wasps (Hymenoptera,Vespidae)

The capacity to recognise a conspecific intruder was investigated in Parischnogaster jacobsoni, Liostenogaster flavolineata and L. vechti, three species of primitively social wasps of the subfamily Stenogastrinae. Results of behavioural experiments carried out in the field showed that females of all three species react pacifically if presented with female nestmates, but aggressively reject an intruder from a conspecific colony. As L. flavolineata and L. vechti both build large clusters of nests, often very close to each other, the recognition capacity among females from different nests, but in the same conspecific cluster, was also investigated. Females of both species were more aggressive towards females from a different colony in the same cluster than towards their female nestmates. Additional experiments on L. flavolineata showed that there was no difference in reaction towards females from colonies nearer or further from the tested colony but within the same cluster, nor towards females from a different cluster. The capacity to recognise an alien conspecific nest containing immature brood was investigated in P. jacobsoni. Adult females of this species, invited to land on an alien nest which had experimentally been exchanged for their own, accepted the new nest and partially destroyed the immature brood. The behaviour of the females when they land on an alien nest, however, suggests that they do recognise the nest as foreign. Acceptance of foreign nests coupled with low immature brood destruction is probably due to the high energetic costs of egg-deposition and larval rearing in stenogastrine wasps. These results suggest that nestmate recognition in these wasps is very efficient, even though they belong to the most primitive subfamily of social wasps. Received: 16 April 1996/Accepted after revision: 9 August 1996     

Trading-off risks from predators and from aggressive hosts

Some birds exploit the aggressiveness of others that are more capable of defending their nests against predators. Usually this behaviour is selected for through improved breeding success for the timid (associate) species. The risk of predation from the more aggressive (host) species is generally rare but may select against the behaviour. The breeding success of red-breasted geese (Branta ruficollis) nesting in association with peregrine falcons (Falco peregrinus) in arctic Siberia was investigated from 1995 to 1999. Nest-predation risk for geese within colonies was positively correlated with distance from the falcons' eyries and negatively correlated with colony size. Though predation risk from the hosts was low, the risk of being attacked by the falcons was high, which we suggest explains why the likelihood of geese deserting their nests was negatively correlated with distance from the eyries. The optimal distance geese nested from eyries was 46 m, though only 37% of pairs managed to nest within 20 m of this. In years with low predation pressure, some geese within colonies nested farther from the falcons. This may be the first evidence to suggest a trade-off between costs and benefits in a nesting association, both of which are directly mediated by the host's nest defence behaviour.     

Blue eggs do not reduce nest predation in the song thrush,Turdus philomelos

Summary Many passerine birds with open cup-shaped nests lay blue or blue-green eggs. In thrushes, blue eggs may be cryptic and provide camouflage by imitating spots of light on green leaves. Alternatively, egg coloration may be selectively neutral because nest predators detect nests and not eggs, or it may be maladaptive because organisms are not always well adapted to their present environment. I evaluated these hypotheses by studying predation on artificial song thrush (Turdus philomelos) nests with quail eggs, painted either white, blue, or spotted (cryptic to a human eye). Corvids were the major nest predators. For concealed as well as exposed nests, I found no differences in the predation rates of nests with white, blue, or spotted eggs. Predators apparently detected the nests, and not the eggs, first. In a second experiment, I placed egg groups without nests in trees to study the effect of color per se. The predation rate of the spotted egg groups was significantly lower than that of the white and blue egg groups, for concealed as well as exposed egg groups. These results suggest that blue eggs in the song thrush are not cryptic but may be selectively neutral or even maladaptive with regard to nest predation.     

Colony fission affects kinship in a social insect

Establishment of new groups is an important step in the life history of a social species. Fissioning is a common mode not only in group proliferation, for instance, as a regular part of the life cycle in the honey bee, but also when multiple females reproduce in the same group, as in multiple-queen ant societies. We studied the genetic consequences of fissioning in the ant Proformica longiseta, based on DNA microsatellites. In P. longiseta, new nests arise by fissioning from the old ones when they grow large, and the daughter nests consist of workers and queens or queen pupae but never both. Our results show that fissioning is not entirely random with respect to kinship. Workers tend to segregate along kin lines, but only when the initial relatedness in the parental nests is low. Workers in a daughter nest also tend to be associated with closely related adult queens, whereas such an association is not detected between workers and queen pupae. Most queens and workers are carried to the daughter nest by a specialized group of transporting workers, suggesting active kin discrimination by them. Fissioning pattern in P. longiseta is different from that found in other social insects with regular fission (e.g., the honey bee, swarm-founding wasps), where no fissioning along kin lines has been found. It does, however, resemble fissioning in another group of social animals: primates.     

Nesting success of rufous-naped wrens (Campylorhynchus rufinucha) is greater near wasp nests

Summary Rufous-naped wrens (Campylorhynchus rufinucha) in northwestern Costa Rica build breeding nests most frequently in ant-acacia trees (Acacia collinsii) and occasionally near wasp nests in ant-acacia trees. By moving occupied wasp nests (Polybia rejecta) to randomly chosen ant-acacia trees with wren nests, I tested the hypothesis that wrens nesting near wasp nests were more likely to fledge young than wrens not nesting near wasp nests. Wrens whose nests were near experimentally relocated wasp nests were significantly more likely to fledge young (37.5% of 16 attempts in 1987 and 75% of 12 attempts in 1988) than were wrens whose nests had no wasp nests placed near them (0% of 16 attempts in 1987 and 20% of 15 attempts in 1988). In 15 cases, repeated nesting attempts occurred in the same trees both with and without experimentally-placed wasp nests. Analysis of these data allowed a comparison of the effect of wasp nests on fledging success while differences among trees were controlled. Within the same tree, nesting attempts associated with wasp nests were significantly more likely to fledge young than nesting attempts without wasp nests. Predation was the primary cause of nest failure, and within forest, white-faced monkeys (Cebus capucinus) were the most important predators. The difference in success of wren nests with and without wasps and observations of predators indicate that enhancement of fledging success was due to deterrence of predatory vertebrates by wasps.     

Is big the best? Queen size,usurpation and nest closure in a primitively eusocial sweat bee (<Emphasis Type="Italic">Lasioglossum malachurum</Emphasis>)

For primitively eusocial insects in which a single foundress establishes a nest at the start of the colony cycle, the solitary provisioning phase before first worker emergence represents a risky period when other, nestless foundresses may attempt to usurp the nest. In the primitively eusocial sweat bee Lasioglossum malachurum (Hymenoptera, Halictidae), spring foundresses compete for nests which are dug into hard soil. Nest-searching foundresses (‘floaters’) frequently inspected nests during this solitary phase and thereby exerted a usurpation pressure on resident queens. Usurpation has been hypothesised to increase across the solitary provisioning phase and favour closure of nests at an aggregation, marking the termination of the solitary provisioning phase by foundresses, before worker emergence. However, our experimental and observational data suggest that usurpation pressure may remain constant or even decrease across the solitary provisioning phase and therefore cannot explain nest closure before first worker emergence. Levels of aggression during encounters between residents and floaters were surprisingly low (9% of encounters across 2 years), and the outcome of confrontations was in favour of residents (resident maintains residency in 94% of encounters across 2 years). Residents were significantly larger than floaters. However, the relationship between queen size and offspring production, though positive, was not statistically significant. Size therefore seems to confer a considerable advantage to a queen during the solitary provisioning phase in terms of nest residency, but its importance in terms of worker production appears marginal. Factors other than intraspecific usurpation need to be invoked to explain the break in provisioning activity of a foundress before first worker emergence.     

Nestmate discrimination in social wasps (Polistes metricus,Hymenoptera: Vespidae)

Summary Laboratory-overwintered paper wasp (Polistes metricus) females preferentially associate on new nests with former nestmates (presumably their sisters) rather than with non-nestmates in the absence of cues associated with their natal nests or nest sites. Females isolated from conspecifics and nests for 74–99 days retain the ability to discriminate nestmates from nonnestmates.     

Physical variability among nest foundresses in the polygynous social wasp,Polistes annularis

Summary An individual's rank in a dominance hierarchy is often based on size or weight, especially in insects. Foundresses of the social wasp, Polistes annularis, vary greatly in size as measured by wing length, dry weight, fat weight, and residue weight after fat has been extracted. Females that emerged from the same nest are much more similar in size to each other than they are to females that emerged from other nests. Within nests however queens are usually larger than their subordinates. Queens that emerged from one nest may be smaller than subordinates that emerged from another nest. We found no evidence of a group of females that are forced into being subordinates because of inadequate feeding as larvae. Females are also probably not attempting to begin new nests with females as different in size as possible from themselves since means and variances of winglength of females on new nests do not differ from those of all females emerging from that natal nest.