Nest defense behavior against ant attacks in post-emergent colonies of wasp Mischocyttarus cerberus (Hymenoptera, Vespidae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

NEST REUSE AND COMMUNAL NESTING IN MICROTHURGE-CORUMBAE (HYMENOPTERA, MEGACHILIDAE), WITH SPECIAL REFERENCE TO NEST DEFENSE

Observations on the nesting activities of Microthurge corumbae, carried out at the University Campus of Ribeirao Preto, São Paulo, Brazil, from 1977 to 1981, indicated that 61.9% of nests were re-used by succeeding generations. Re-use by one generation was more frequent than by two generations, and re-use by a third was observed only once. Nests were re-used by one or several females. Single females were more frequently in the first re-use. In these cases nest re-use did not differ essentially from the solitary foundation of a new nest, except for the adoption of a pre-existing nest without excavation. In multifemale nests, analysis of relative age (wing wear), ovarian and spermathecal conditions of associated females and the content of nests at excavation indicated that the social pattern in such colonies is communal. There is some evidence that the associated females are relatives. The chalcidoid wasp Leucospis was the principal nest parasite, and ants of the genus Crematogaster were nest predators. In multifemale nests, the rate of parasitism was significantly lower than in solitary nests, indicating that nest-sharing resulted in improved nest defense. on the other hand. The absence of predation on immatures of the first generation of M. corumbue in multifemale nests suggests that such nests are also more resistant to attack by predators.

Nest defense behavior against the attack of ants in colonies of pre-emergent Mischocyttarus cerberus (hymenoptera, vespidae)

The study was carried out at the UNESP Rio Claro campus (SP), where biotests consisting of simulated ant attacks were performed in colonies of Mischocyttarus cerberus. The behaviors of the wasps were recorded with a camcorder, for further analysis. This analysis was done using the Mann-Whitney U test and the Principal Component Analysis. In the pre-emergence development stage, colonies with a single foundress defend the nest only after the first larvae appear. When there are only eggs in the nest, the wasp abandons the nest. Before leaving, the wasp rubs its gaster against the nest, releasing the ant repellent secretion. When the nest contains larvae or larvae and pupae, the foundress defends the colony, vibrating its wings, pumping her abdomen and biting the ant.

Male behavior of Mischocyttarus cerberus (Hymenoptera, Vespidae) during attacks on the nest by ants

The objectives of the present study were to verify if the males of the wasp Mischocyttarus cerberus participate in defense of the nest as they react to attack by ants, and to observe if they use chemical defense. First, wasps were stimulated with an empty pair of tweezers and a second stimulation was done with a pair of tweezers containing an ant, Camponotus crassus. The behaviors of the wasps were recorded and analyzed. The males can participate in defense of the nest, presenting some aggressive behaviors, even so, most of the time they ignore the stimulation, hide behind the nest and stay immobile.

Nest stolen: the first observation of nest predation by an invasive exotic marmoset (Callithrix penicillata) in an agricultural mosaic

Invasive exotic species can negatively impact local biodiversity. We present here a report of a nest predation of an endemic bird species, variable oriole (Icterus pyrrhopterus) by the introduced black-tufted marmoset (Callithrix penicillata)in an agricultural landscape highly disturbed by human activities. Two nestlings were predated, by adults of the introduced marmoset during two alternate days. Antipredator behavior and vocal mimicry were observed in variable oriole, while copulation was observed in black-tufted marmoset during the predation. The use of mobbing against predators by I. pyrrhopterus was observed and it is described here by the first time. The potential impact of the introduced marmosets to local biodiversity is discussed.

A morphologically specialized soldier caste improves colony defense in a neotropical eusocial bee

Division of labor among workers is common in insect societies and is thought to be important in their ecological success. In most species, division of labor is based on age (temporal castes), but workers in some ants and termites show morphological specialization for particular tasks (physical castes). Large-headed soldier ants and termites are well-known examples of this specialization. However, until now there has been no equivalent example of physical worker subcastes in social bees or wasps. Here we provide evidence for a physical soldier subcaste in a bee. In the neotropical stingless bee Tetragonisca angustula, nest defense is performed by two groups of guards, one hovering near the nest entrance and the other standing on the wax entrance tube. We show that both types of guards are 30% heavier than foragers and of different shape; foragers have relatively larger heads, whereas guards have larger legs. Low variation within each subcaste results in negligible size overlap between guards and foragers, further indicating that they are distinct physical castes. In addition, workers that remove garbage from the nest are of intermediate size, suggesting that they might represent another unrecognized caste. Guards or soldiers are reared in low but sufficient numbers (1-2% of emerging workers), considering that <1% usually perform this task. When challenged by the obligate robber bee Lestrimelitta limao, an important natural enemy, larger workers were able to fight for longer before being defeated by the much larger robber. This discovery opens up opportunities for the comparative study of physical castes in social insects, including the question of why soldiers appear to be so much rarer in bees than in ants or termites.

Reduction in Predator Defense in the Presence of Neighbors in a Colonial Fish

Predation pressure has long been considered a leading explanation of colonies, where close neighbors may reduce predation via dilution, alarming or group predator attacks. Attacking predators may be costly in terms of energy and survival, leading to the question of how neighbors contribute to predator deterrence in relationship to each other. Two hypotheses explaining the relative efforts made by neighbors are byproduct-mutualism, which occurs when breeders inadvertently attack predators by defending their nests, and reciprocity, which occurs when breeders deliberately exchange predator defense efforts with neighbors. Most studies investigating group nest defense have been performed with birds. However, colonial fish may constitute a more practical model system for an experimental approach because of the greater ability of researchers to manipulate their environment. We investigated in the colonial fish, Neolamprologus caudopunctatus, whether prospecting pairs preferred to breed near conspecifics or solitarily, and how breeders invested in anti-predator defense in relation to neighbors. In a simple choice test, prospecting pairs selected breeding sites close to neighbors versus a solitary site. Predators were then sequentially presented to the newly established test pairs, the previously established stimulus pairs or in between the two pairs. Test pairs attacked the predator eight times more frequently when they were presented on their non-neighbor side compared to between the two breeding sites, where stimulus pairs maintained high attack rates. Thus, by joining an established pair, test pairs were able to reduce their anti-predator efforts near neighbors, at no apparent cost to the stimulus pairs. These findings are unlikely to be explained by reciprocity or byproduct-mutualism. Our results instead suggest a commensal relationship in which new pairs exploit the high anti-predator efforts of established pairs, which invest similarly with or without neighbors. Further studies are needed to determine the scope of commensalism as an anti-predator strategy in colonial animals.

Reticence or vigilance at the nest: a cruel bind for the endangered Black-capped Vireo

Breeding birds vocalize to find mates and establish and defend territories, but these same critical communications may also attract predators or brood parasites, placing birds in a cruel bind. Although vigilant birds may better maintain social relationships with mates and neighbors through frequent vocalizations, reticent birds may reduce risk to their nests by being relatively quiet and making infrequent vocalizations. Selection for vocalization patterns that minimize brood parasitism might be particularly strong for birds that are unable to fledge both their own young and the parasite. Temporal plasticity in the frequency of vocalizations near nests, however, may allow birds to balance trade-offs and optimize nest-defense strategies. The Black-capped Vireo (Vireo atricapilla) is an endangered songbird that faces intensive brood parasitism in areas where Brown-headed Cowbirds (Molothrus ater) are present. Vireo nests that produce cowbird fledglings always fail to fledge vireo young. We recorded vocalizations at vireo nests across three nesting stages (building, laying, and early incubation) and three periods of the day (morning, midday, and evening) and compared vocalization frequency with eventual depredation or parasitism fate as well as local cowbird density to test two hypotheses. The predator-attraction hypothesis predicts that predators will be attracted by frequent vocalizations, whereas cowbirds will parasitize nests with relatively quiet parents and less predation risk; thus, vireos will experience trade-offs between reticence and vigilance in mediating specific risks. The parasite-assessment hypothesis predicts that vireos will become more secretive as local cowbird densities increase. Vireo vocalization response to nest predation and parasitism risk interacted with nest stage, and we found little evidence of risk mediation through vocalizations except during the building stage. Vireos, however, did benefit overall by optimizing temporal patterns in vocalizations. Vireo nests were less likely to be depredated or parasitized if males vocalized most during laying and least during the middle of the day. Birds vocalized more during the midday and less during the laying period when local cowbird densities were higher, however, perhaps demonstrating limited plasticity in social communication.

Living on voles - plastic life of the Ural owl

Individuals face variable environmental conditions during their life. This may be due to migration, dispersion, environmental changes or, for example, annual variation in weather conditions. Genetic adaptation to a novel environment happens through natural selection. Phenotypic plasticity allows, however, a quick individual response to a new environment. Phenotypic plasticity may also be beneficial for individual if the environment is highly variable. For example, eggs are costly to produce. If the food conditions vary significantly between breeding seasons it is useful to be able to adjust the clutch and egg size according to the food abundance. In this thesis I use Ural owl vole system to study phenotypic plasticity and natural selection using a number of reproduction related traits. The Ural owl (Strix uralensis) is a long-lived and sedentary species. The reproduction and survival of the Ural owl, in fact their whole life, is tied to the dramatically fluctuating vole densities. Ural owls do not cause vole cycles but they have to adjust their behaviour to the rather predictable population fluctuations of these small mammals. Earlier work with this system has shown that Ural owl laying date and clutch size are plastic in relation to vole abundance. Further, individual laying date clutch size reaction norms have been shown to vary in the amount of plasticity. My work extends the knowledge of natural selection and phenotypic plasticity in traits related to reproduction. I show that egg size, timing of the onset of incubation and nest defense aggressiveness are plastic traits with fitness consequences for the Ural owl. Although egg size is in general thought to be a fixed characteristic of an individual, this highly heritable trait in the Ural owl is also remarkably plastic in relation to the changes in vole numbers, Ural owls are laying the largest eggs when their prey is most abundant. Timing of the onset of incubation is an individual-specific property and plastic in relation to clutch size. Timing of incubation is an important underlying cause for asynchronous hatching in birds. Asynchronous hatching is beneficial to offspring survival in Ural owl. Hence, timing of the onset of incubation may also be under natural selection. Ural owl females also adjust their nest defense aggressiveness according to the vole dynamics, being most aggressive in years when they produce the largest broods. Individual females show different levels of nest defense aggressiveness. Aggressiveness is positively correlated with the phenotypic plasticity of aggressiveness. As elevated nest defense aggressiveness is selected for, it may promote the plasticity of aggressive nest defense behaviour. All the studied traits are repeatable or heritable on individual level, and their expression is either directly or indirectly sensitive to changes in vole numbers. My work considers a number of important fitness-related traits showing phenotypic plasticity in all of them. Further, in two chapters I show that there is individual variation in the amount of plasticity exhibited. These findings on plasticity in reproduction related traits suggest that variable environments indeed promote plasticity.

Breeding of the Rufous-thighed Hawk (Accipiter erythronemius) in Argentina and Brazil

The Rufous-thighed, Hawk (Accipiter erythronemius) is a poorly-known, small raptor that dwells in forests and wooded savannas in south and eastern Brazil, Taraguay, eastern Bolivia, north and central Argentina, and Uruguay. We studied breeding biology and food habits of the Rufous-thighed Hawk from 1994 to 2007. We documented 14 breeding attempts recorded at nine nest sites in eight localities in Argentina and Brazil. Rufous-thighed Hawks were year-round residents at the study areas. The breeding season was c. 38 weeks long (June through March). Territorial behavior and courtship spanned for 22 weeks starting in mid June. Nest-building started in August. Nestlings were observed throughout December and into early January while fledglings were observed from late December through late March. Most nest sites were conifer plantations averaging 32.2 (SD = 10) years old, 12.6 ha (SD = 11.3) in size with trees averaging d 37.5 cm (SD = 13.5) of diameter at breast height. Nests were placed 20.2 m high (SD = 5.7), near the trunks of non-emergent conifers (Pinus sp. or Araucaria angustifolia). Nests were made of sticks, greater diameter averaging 48.8 cm (SD = 11.4), while nest depth averaged 23.3 cm (SD = 14). Prey items were birds (n = 49) ranging in size from c. 10 g [House Wren (Troglodytes, aedon)] to c. 140 g [White-tipped Dove (Leptotila verreauxi)]. Young started to chase birds at the age of 42-45 days in mid January. Adult males were primarily food providers but also aided in nest defense against other raptors. Females incubated, brooded, and fed young, and took leading roles in defense against humans. Rufous-thighed Hawks had unlined nests placed high in trees, and breeding season was markedly protracted.

A resource range invariance rule for optimal offspring size predicts patterns of variability in parental phenotypes

Previous analysis of the rules regarding how much more a female should invest in a litter of size C rather than producing a litter with one more offspring revealed an invariance relationship between litter size and the range of resources per offspring in any litter size. The rule is that the range of resources per offspring should be inversely proportional to litter size. Here we present a modification of this rule that relates litter size to the total resources devoted to reproduction at that litter size. The result is that the range of resources devoted to reproduction should be the same for all litter sizes. When parental phenotypes covary linearly with resources devoted to reproduction, then those traits should also show equal ranges within each litter size category (except for litters of one). We tested this prediction by examining the range in body size (=total length) of female mosquito fish (Gambusia hubbsi) at different litter sizes. Because resources devoted to reproduction may take many forms (e.g., nest defense), this prediction may have broad applicability.

Division of labour within cooperatively breeding groups

Within cooperative societies, group members share in caring for offspring. Although division of labour among group members has been relatively well studied in insects, less is known about vertebrates. Most studies of avian helping focus solely on the extent to which helpers provision the offspring, however, helpers can participate in everything from nest building to predator defence. Bad provisioners may, for example, not be as 'uncooperative' as they appear. if they are good defenders. Thus, the distribution of helping tasks between group members should have important implications for our interpretation of group dynamics. Here, we compare two distinct forms of helping behaviour in the cooperatively breeding noisy miner (Manorina melanocephala): chick provisioning and mobbing nest predators. We show that the way in which individual helpers invest in these two helping behaviours varies enormously across individuals and among social groups. Good provisioners often contributed relatively little to mobbing and vice versa. Indeed, (18%) of helpers only mobbed, 22% just provisioned, whereas 60% of helpers performed both forms of helping. Across nests, provisioning was significantly negatively correlated with mobbing effort. We suggest that small differences in the costs or benefits of different aspects of helping (due to differences in age, relatedness or social status) have a big impact on the division of labour within a group. Consequently, social groups can be made up from individuals who often specialise in one helping behaviour, and/or helpers who perform a number of behaviours to differing degrees. Division of labour within social groups will, therefore, have important consequences for the maintenance of cooperatively breeding in vertebrates.

Colony defense behavior of the primitively eusocial wasp, Mischocyttarus cerberus is related to age

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)