Evaluation of the Predatory Wasp, Ancistrocerus gazella, for Biological Control of Leafrollers in Otago Fruit Crops: 1. Prey Composition, Nest Structure and Wasp Productivity from Artificial Nests

Groups of artificial nest tubes were deployed in the field for colonization by the accidentallyimported wasp, Ancistrocerus gazella (Panzer), a reported important predator of lepidopterous larvae, especially pest tortricids. There were 3-4 nest sites in each of seven study areas, and four tube lengths (75, 100, 125, 150 mm) were compared. Nests were dissected at various stages after occupancy to determine the prey composition and nest structure, and the numbers and sex of adults produced. The wasps collected larvae of nine lepidopterous families, including many tortricids, but pest leafrollers comprised only 4.1% and 1.5% of all larvae collected respectively in 1995-96 and 1996-97. Prey included beneficial species feeding on weeds and the larvae of some non-pest native insects. Prey species composition was highly site-dependent and was a major determinant of the numbers of larvae per cell. Cells for male wasps were smaller and contained fewer larvae than those for females. Tube length did not affect colonization rates but more cells and adult wasps were produced in longer tubes. The results cast serious doubt on the proposed management of A. gazella for biological control of leafrollers, but suggest that this approach could be useful for other pests, such as Coleophora spp., which were collected in large numbers.     

The Weaver Wasp: Spinning Fungus into a Nest

Wasp nests range from simple to complex structures made of paper or mud. Here, we show that a Neotropical wasp of the genus Nitela builds its nest entirely by weaving endophytic fungal hyphae and spider silk harvested from the leaves growing in the understory of the rain forest in French Guiana.     

Males Exhibit Novel Relationships of Dominance with Nest Departure in the Social Paper Wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

In many animal societies, an individual's opportunities to reproduce correspond to their position in a dominance hierarchy. In primitively eusocial paper wasps, female social dominance and direct reproduction are positively associated with taking food from nest mates, but both negatively correlate with foraging flights. Female paper wasps reproduce by laying eggs in their nest, while males reproduce by leaving the nest to copulate. Males of Mischocyttarus mastigophorus are unusual because they reside on their natal nests for prolonged periods, and males direct aggression towards female nestmates. Mischocyttarus mastigophorus males provide an opportunity to further analyse the relationships between dominance, nutrition and nest departure. We found males' rates of social aggression, food taking behaviour, and rates of nest departure were positively correlated. Furthermore, males that initiated aggression at younger ages also began flying earlier, and males engaged in all three behaviours at younger ages than females. Male social aggression may lead to nutrient acquisition, which enable males to depart from the nest. We hypothesise that male nest departure could be adaptive if it increases male mating opportunities. This could account for the positive relationship between dominance and nest departure for males.     

Genetic Relatedness of Low Solitary Nests of Apis dorsata from Marang, Terengganu, Malaysia

Knowledge on the population of genetic structure and ecological behaviour of Apis dorsata from Peninsular Malaysia is needed for effective management and conservation of this species since unsustainable whole solitary low nest cutting for product harvesting is the current common practice here. The analysis of 15 single locus DNA microsatellite markers on samples from 20 solitary nests of A. dorsata showed that while these markers were polymorphic, high intracolonial relatedness existed. Furthermore, in general, slightly negative values of intercolony relatedness (R) among the nests of A. dorsata were found. However, positive values of mean intercolony relatedness were observed between 54 pairs of nests out of 190 possible combinations. The R values among nest pairs 3-4 and 3-5 was higher than 0.50 showing that their queens were half siblings, whereas nest pair 19-20 showed relatedness of 0.95 indicating that the same queen was sampled. The results that we obtained could not conclusively support the hypothesis of this study that the honey hunters in Marang district of Malaysia repeatedly harvest the same nest located at a different site and at a different time during the same honey harvesting season. However, our finding of an appreciable level of intercolonial relatedness between several pairs of nests in this pioneer study indicated that a comprehensive study with a larger sample size of solitary nests found throughout the region would be necessary to provide concrete proof for this novel idea.     

Poor Odors, Strength, and Persistence Give Their Rewards to Mutilla europaea Visiting Dangerous Wasp Nests

Social insect colonies are attractive for many arthropods. The rare velvet ant, Mutilla europaea, visit colonies of Polistes wasps, but to date it was unclear which resources it targeted therein. Our field observations and bioassays showed that velvet ants visit the colonies of the social wasp Polistes biglumis almost undisturbed and may feed on larval wasp saliva. Chemical insignificance and resistance are the characteristics that allow velvet ants to visit unharmed wasp colonies and gain such a nutritious reward.     

Age,but not Sex or Genetic Relatedness,Shapes Raccoon Dominance Patterns

Raccoons are generally regarded as solitary, yet several studies have found that raccoons frequently form social affiliations. One benefit to sociality in many mammal species is that relatives and close associates can form coalitions against third parties during agonistic encounters. We tested whether raccoon dominance patterns were influenced by age, sex, genetic relatedness, and association patterns at two anthropogenic feeding stations in an urban forest. We found that genetic relatedness had no significant effect on patterns of agonism at one of the feeding stations. At the second feeding station, raccoons were more likely to act aggressively toward close relatives, which is opposite of the predicted pattern. However, when we controlled for the number of times raccoons arrived at feeding stations in close proximity, the effect of relatedness on dominance patterns was not significant at either feeding station. These results suggest that relatedness plays little or no role in shaping dominance patterns of raccoons. Older raccoons were ranked significantly higher in the dominance hierarchies regardless of sex. This pattern leads us to conclude that age is the primary factor driving the outcome of aggressive interactions in raccoons at our study site. Despite frequent social interactions at the study site, the patterns of raccoon dominance more closely resemble patterns found in solitary animals. To confirm the generality of these results and to better understand the evolution of raccoon social behavior, similar studies need to be undertaken in other raccoon populations.     

Effects of Relatedness,Dominance, Age,and Association on Reciprocal Allogrooming by Captive Impala

Wild impala display a highly reciprocal allogrooming system that, by virtue of its frequency and high degree of reciprocity, is unique among ungulates. A herd of 35 free-ranging captive impala provided opportunity to examine the degree of reciprocity of allogrooming exchanges and the influence of relatedness, dominance, age and association on partner preferences and distribution of grooming between allogrooming partners. As in wild impala, the exchange of allogrooming bouts in the captive impala was highly reciprocal regardless of partners. Kinship and dominance had no influence on partner preference or distribution of grooming between partners. Although mothers showed a significant preference to allogroom with their unweaned offspring, this preference practically disappeared with older offspring. Age-mates (no greater than 6 mo apart) tended to associate with one another and spatial proximity was positively correlated with grooming partner preference. It was not clear whether impala actively sought out age-mates for grooming, or randomly chose grooming partners from nearby age-mates. The failure to find a role for kinship and dominance is counter to what has generally been found in most Old World terrestrial primate studies. The absence of pronounced social influences, coupled with the known effectiveness of grooming in removing ectoparasites, suggest that a utilitarian role, especially removal of ticks, is an important function of the impala reciprocal allogrooming system.     

Nest Predation Risk on Ground and Shrub Nests in Forest Margin Areas of Sulawesi, Indonesia

Forest loss and fragmentation in Indonesia may seriously affect the survivorship of forest birds and lead to local extinction of bird populations. We used 786 artificial nests baited with quail eggs to examine the effect of habitat alteration on nest predation in Lore Lindu National Park, Sulawesi. Natural forest and four habitats of forest margin areas: forest edge, forest gardens, coffee plantations, and secondary forest, were studied. Two types of artificial nests, ground and shrub nests were placed in these habitats at two different locations for a period of 8 days. In addition, we used automatic cameras and cage-traps to identify the predators. Nests in shrubs experienced significantly higher predation rates in forest margin areas than in natural forest. Predation on ground nests did not differ significantly between these habitat types, but was significantly higher than that on shrub nests in each habitat except forest edge. Rodents were the most common predators of both nests, but shrub nests were also susceptible to Dwarf cuscus (Strigocuscus celebensis), squirrels, and tree snakes. The nest predation rates we found were among the highest found in tropical rainforests, probably a consequence of the unique predator assemblages of Sulawesi. These results suggest that egg survival is negatively affected by human intervention and that human-induced habitats might have only limited importance for the conservation of Sulawesi's largely endemic understorey avifauna. These considerations might be important since forest margins comprise significant proportions of protected areas on Sulawesi and play an important role in future Park zoning concepts as well as in conservation-oriented land use management.     

The Function of Male Dominance in the Eusocial Wasp, Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Males of a Neotropical eusocial wasp, Mischocyttarus mastigophorus , are dominant over their female nest mates. Mischocyttarus mastigophorus males behave aggressively toward females, while females rarely bite or chase males. Aggressive interactions between the sexes are behaviorally indistinguishable from dominance interactions among females. Males are long-lived as adults, and can reside on nests for periods of at least one month. Furthermore, males comprise a large proportion of post-emergence colony populations throughout much of the colony cycle. Males on the nest perform maintenance tasks at low rates, but contribute little other labor to their colonies. Males do not forage, but consume a disproportionate amount of the food (nectar and insect prey) collected by workers. Males in some colonies direct disproportionate amounts of aggression toward their queens, which may further contribute to males' food procurement. These data suggest that adult males represent a considerable energetic and labor cost to their colonies. I hypothesize that the dominance structure of M. mastigophorus directs food resources to adult males, with the function of increasing their longevity. Increased male longevity may be selectively advantageous in tropical species such as M. mastigophorus that found new colonies throughout much or all of the year. When females initiate new nests over much of the year, individual males' mating opportunities may be temporally distributed, favoring longer adult lifespans. Male dominance is predicted to occur in other populations of independent-founding Neotropical Polistinae with asynchronous colony foundation.     

Behavioral Profiles Related to Dominance Hierarchy in Associated Foundresses of Belonogaster juncea juncea (F.) (Hymenoptera: Vespidae)

This study was conducted to examine the phenomenon of task partitioning among associations of foundresses of the primitively eusocial wasp Belonogaster juncea juncea (F.). The time–activity budget for five main behavioral categories (foraging, building, feeding, inactivity, and reproduction) for each foundress belonging to trigynic associations was recorded using the instantaneous scanning technique. After ranking the individuals on the basis of agonistic encounters, data were submitted to a canonical discriminant analysis. The results show behavioral differences between individuals of each rank. The proportion of time allocated to foraging behavior is a good rank index. The females of the first rank spent less time on foraging behavior and significantly more time (13.6% of their time) on reproductive behavior than females of other ranks. The females of the second rank also spent less time on foraging behavior and showed a tendency toward building behavior. The females of the third rank spent significantly more time (77.28% of their time) on foraging behavior. The behavioral profile of each foundress was therefore determined by its rank on the dominance scale.     

Effects of Edge Habitat and Nest Characteristics on Depredation of Artificial Nests in Fragmented Australian Tropical Rainforest

Variation in nest predation levels associated with rainforest fragmentation (edge effects) was assessed in Australia's Wet Tropics bioregion. Artificial nests were placed in the forest understorey at seven edge sites where continuous forest adjoined pasture, seven interiors (about 1 km from the edge), and six linear riparian forest remnants (50–100 m wide) that were connected to continuous forest. Four nest types were also compared, representing different combinations of two factors; height (ground, shrub) and shape (open, domed). At each site, four nests of each type, containing one quail egg and two model plasticine eggs, were interspersed about 15 m apart within a 160 m transect during September–October 2001. Predators were identified from marks on the plasticine eggs. The overall depredation rate was 66.5% of 320 nests' contents damaged over a three-day period. Large rodents, especially the rat Uromys caudimaculatus, and birds, especially the spotted catbird Ailuroedus melanotis, were the main predators. Mammals comprised 56.5% and birds 31.0% of predators, with 12.5% of unknown identity. The depredation rate did not vary among site-types, or between open and domed nests, and there were no statistically significant interactions. Nest height strongly affected depredation rates by particular types of predator; depredation rates by mammals were highest at ground nests, whereas attacks by birds were most frequent at shrub nests. These effects counterbalanced so that overall there was little net effect of nest height. Mammals accounted for 78.4% of depredated ground nests and birds for at least 47.4% of shrub nests (and possibly up to 70.1%). The main predators were species characteristic of rainforest, rather than habitat generalists, open-country or edge specialists. For birds that nest in the tropical rainforest understorey of the study region, it is unlikely that edges and linear remnants presently function as ecological population sinks due to mortality associated with increased nest predation.     

Social Network Analysis of Male Dominance in the Paper Wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Social aggression is a pervasive feature of insect societies. In eusocial Hymenoptera, aggression among females can affect task performance and competition over direct reproduction (egg laying); in most species males participate in social interactions relatively rarely. Males of the independent-founding paper wasp Mischocyttarus mastigophorus are exceptional: they are aggressive toward female nestmates, leading us to explore the function of this unusual behavior. We applied social network analyses to data on M. mastigophorus social aggression to quantify sex differences in giving and receiving social aggression. The network analyses supported the pattern of biased male aggression toward female nestmates; females are relatively rarely aggressive to males. We then asked whether male aggression toward females was biased by females’ relative ovary development. Males were more aggressive toward females with better-developed ovaries, opposite to patterns of aggression among females. Because food brought to the colonies is often monopolized by dominant females, we suggest that males direct aggression toward socially dominant females with better-developed ovaries to obtain food. The implications of biased male aggression for female task performance and physiology are unknown.

     

Queens of the Paper Wasp Polistes fuscatus (Hymenoptera: Vespidae) Discriminate among Larvae on the Basis of Relatedness

The ability of queens of the paper wasp Polistes fuscatus to recognize larvae on the basis of colony origin was investigated in the laboratory. In a blind, binary choice paradigm, queens discriminated between larvae from their own colony and unrelated larvae, and between larvae from a sister colony and unrelated larvae. However, queens failed to discriminate between larvae from their own colony and larvae from a sister colony, indicating that queens of multiple-foundress colonies may lack the ability to make intracolonial brood discriminations on the basis of relatedness. These results demonstrate that recognition of larvae is mediated by larval-borne, genetically specified odors. Gas chromatograph/mass spectral analysis revealed that adults and larvae have similar cuticular hydrocarbon profiles, suggesting that the cues mediating larval and adult recognition may be the same.     

Habitat Quality, Social Dominance and Dawn Chorus Song Output in Black-Capped Chickadees

Individuals that settle in poor habitats may reduce resource investments in various life history traits; for example, resources may be withheld from costly advertisement signals. There may be geographic variation in advertisement levels that correspond with habitat quality; however, this is poorly documented and it is unclear whether such habitat effects have consequences for the function of mate‐choice signals within habitats. We examined song output of male black‐capped chickadees (Poecile atricapillus) during the dawn chorus in two contrasting habitats (mature forest vs. young forest) known to differ in quality. Dominance rank is indicative of phenotypic quality in this species and was assessed during the preceding winter. We measured the song output of males participating in the dawn chorus during the nesting/egg laying period. Males living in young forest had reduced song output; however, a habitat–rank interaction term showed that dominant birds accounted for most of this difference. This suggests that signal reliability might be lost in poor habitats. We generate several hypotheses to explain these findings.     

Florida Harvester Ant Nest Architecture,Nest Relocation and Soil Carbon Dioxide Gradients

Colonies of the Florida harvester ant, Pogonomyrmex badius, excavate species-typical subterranean nests up the 3 m deep with characteristic vertical distribution of chamber area/shape, spacing between levels and vertical arrangement of the ants by age and brood stage. Colonies excavate and occupy a new nest about once a year, and doing so requires that they have information about the depth below ground. Careful excavation and mapping of vacated and new nests revealed that there was no significant difference between the old and new nests in any measure of nest size, shape or arrangement. Colonies essentially built a replicate of the just-vacated nest (although details differed), and they did so in less than a week. The reason for nest relocation is not apparent. Tschinkel noted that the vertical distribution of chamber area, worker age and brood type was strongly correlated to the soil carbon dioxide gradient, and proposed that this gradient serves as a template for nest excavation and vertical distribution. To test this hypothesis, the carbon dioxide gradient of colonies that were just beginning to excavate a new nest was eliminated by boring 6 vent holes around the forming nest, allowing the soil CO₂ to diffuse into the atmosphere and eliminating the gradient. Sadly, neither the nest architecture nor the vertical ant distribution of vented nests differed from either unvented control or from their own vacated nest. In a stronger test, workers excavated a new nest under a reversed carbon dioxide gradient (high concentration near the surface, low below). Even under these conditions, the new and old nests did not differ significantly, showing that the soil carbon dioxide gradient does not serve as a template for nest construction or vertical worker distribution. The possible importance of soil CO₂ gradients for soil-dwelling animals is discussed.     

Relatedness,Conflict, and the Evolution of Eusociality

The evolution of sterile worker castes in eusocial insects was a major problem in evolutionary theory until Hamilton developed a method called inclusive fitness. He used it to show that sterile castes could evolve via kin selection, in which a gene for altruistic sterility is favored when the altruism sufficiently benefits relatives carrying the gene. Inclusive fitness theory is well supported empirically and has been applied to many other areas, but a recent paper argued that the general method of inclusive fitness was wrong and advocated an alternative population genetic method. The claim of these authors was bolstered by a new model of the evolution of eusociality with novel conclusions that appeared to overturn some major results from inclusive fitness. Here we report an expanded examination of this kind of model for the evolution of eusociality and show that all three of its apparently novel conclusions are essentially false. Contrary to their claims, genetic relatedness is important and causal, workers are agents that can evolve to be in conflict with the queen, and eusociality is not so difficult to evolve. The misleading conclusions all resulted not from incorrect math but from overgeneralizing from narrow assumptions or parameter values. For example, all of their models implicitly assumed high relatedness, but modifying the model to allow lower relatedness shows that relatedness is essential and causal in the evolution of eusociality. Their modeling strategy, properly applied, actually confirms major insights of inclusive fitness studies of kin selection. This broad agreement of different models shows that social evolution theory, rather than being in turmoil, is supported by multiple theoretical approaches. It also suggests that extensive prior work using inclusive fitness, from microbial interactions to human evolution, should be considered robust unless shown otherwise.     

Relatedness, phylogeny, and evolution of the fungi

Recent advances in fungal systematics are reviewed in relation to our previous studies. The usefulness of the integrated analysis of genotypic (especially 18S rRNA gene sequence comparisons) and phenotypic (especially ultrastructural and chemotaxonomic data) characters has been emphasized for the major groups and selected taxa of the fungi, and the impact to fungal systematics and evolution is discussed. Our noteworthy studies and findings are: 1) polyphyly of the chytridiomycetes and zygomycetes, 2) phylogenetic origin of the entomophthoralean fungi includingBasidiobolus, 3) detection of a major new lineage “Archiascomycetes,” comprisingTaphrina, Protomyces andSaitoella, Schizosaccharomyces, andPneumocystis, within the Ascomycota, and its phylogenetic and evolutionary significance, 4) polyphyletic origins of species in the anamorphic genusGeosmithia, and 5) phylogenetic placement ofMixia osmundae, species correctly and incorrectly assigned to the genusTaphrina, and basidiomyceotus yeasts. The newest 18S rDNA sequence-based neighbor-joining trees of the Ascomycota are demonstrated. “Traditional studies of evolution have amply demonstrated that evolution at the phenotypic level is characterized by adaptation and opportunism, irregularity in pace, and inequality of rates among lineages. In contrast, studies of molecular evolution have revealed quite different features characterized by changes that are conservative in nature, random in pattern (independent of phenotypic characters), and quite regular in pace with equal rates among diverge [sic] lineages for a given protein”. (Kimura, M. 1983. The neutral theory of molecular evolution, pp. 308–309, Cambridge University Press, Cambridge.) Recipient of the 2nd Mycological Society of Japan's Excellent Achievement Award, 1998; the awarding lecture was given at the 42nd Annual Meeting of the Mycological Society of Japan, 16 May, 1998, Kyoto University, Kyoto. This review is based mainly on the publications intended for the Award.     

Functional and Phylogenetic Relatedness in Temporary Wetland Invertebrates: Current Macroecological Patterns and Implications for Future Climatic Change Scenarios

In freshwater ecosystems, species compositions are known to be determined hierarchically by large to small‑scale environmental factors, based on the biological traits of the organisms. However, in ephemeral habitats this heuristic framework remains largely untested. Although temporary wetland faunas are constrained by a local filter (i.e., desiccation), we propose its magnitude may still depend on large-scale climate characteristics. If this is true, climate should be related to the degree of functional and taxonomic relatedness of invertebrate communities inhabiting seasonal wetlands. We tested this hypothesis in two ways. First, based on 52 biological traits for invertebrates, we conducted a case study to explore functional trends among temperate seasonal wetlands differing in the harshness (i.e., dryness) of their dry season. After finding evidence of trait filtering, we addressed whether it could be generalized across a broader climatic scale. To this end, a meta-analysis (225 seasonal wetlands spread across broad climatic categories: Arid, Temperate, and Cold) allowed us to identify whether an equivalent climate-dependent pattern of trait richness was consistent between the Nearctic and the Western Palearctic. Functional overlap of invertebrates increased from mild (i.e., Temperate) to harsher climates (i.e., Arid and Cold), and phylogenetic clustering (using taxonomy as a surrogate) was highest in Arid and lowest in Temperate wetlands. We show that, (i) as has been described in streams, higher relatedness than would be expected by chance is generally observed in seasonal wetland invertebrate communities; and (ii) this relatedness is not constant but climate-dependent, with the climate under which a given seasonal wetland is located determining the functional overlap and the phylogenetic clustering of the community. Finally, using a space-for-time substitution approach we suggest our results may anticipate how the invertebrate biodiversity embedded in these vulnerable and often overlooked ecosystems will be affected by long-term climate change.