Effect of prey quantity and temperature on nest demography of social wasps

Abstract. 1. To determine the effect of prey quantity and temperature on nest demography of social wasps ( Polistes fuscatus ), field experiments were conducted, in which wasps were provided with a low quantity of caterpillars (approximately nest subsistence level) or high quantity (three times as much). In addition, in the third year, the nest boxes were modified to be relatively cool (white and insulated) or warm (black).
2. In 1997 and 1998, high-food nests had a high proportion of cells containing developing offspring, produced more offspring, and had disproportionately more female offspring compared with low-food nests.
3. In 1999, the cool and warm nest boxes exhibited a daily average difference of 1.3 °C and a maximum difference of 6.9 °C. The warm high-food nests produced more offspring than any other treatment. The cool high-food nests produced similar numbers of offspring as the cool low-food and warm low-food treatments.
4. As indicated by a surplus of uncapped cells, foundresses of low-food nests, which would have had the egg-laying potential to produce many offspring, apparently sacrificed eggs to provide a food supplement for the developing offspring. This pattern plus more time observed foraging suggests considerable expenditure of energy by foundresses, in response to the low supply of food.
5. Importantly, these results indicate that an interactive effect of prey quantity and temperature on offspring production occurred, which may reflect the different effects on developmental rate and growth rate at cool versus warm nest temperatures.     

Context-dependent running speed in funnel-web spiders from divergent populations

1.  Locomotor performance can influence individual fitness through several ecological contexts, such as prey capture and predator escape. One means of determining which contexts act as significant selective forces on running speed is to quantify individual speed in each context. The underlying hypothesis is that animals will exhibit their highest speeds in contexts most crucial to fitness.
2.  We measured running speeds in three ecological contexts (prey capture, fleeing predators and territory defence) in lab-reared offspring of the funnel-web spider Agelenopsis aperta collected from two arid grassland and two riparian populations. Arid populations experience little predation pressure, are prey limited, and are highly territorial; riparian populations experience high predation, have high prey availability, and are less territorial in nature.
3.  The offspring of arid individuals exhibited their highest burst speeds in territory defence, and ran more slowly in response to predator threats. The offspring of riparian populations, however, ran fastest when responding to predatory threats and displayed lower velocities in prey capture and territory defence. Thus, our findings support the hypothesis that A. aperta are selected to exhibit their highest speeds in contexts most important to their fitness.
4.  Contextual use of running speed can differ among conspecific populations experiencing differing selective forces on locomotion.     

Development and evolution

Unlike many other animals whose sex ratios have been studied, parasitic wasps are able to determine the sex of their offspring. It is known that parasitic wasps sometimes produce different offspring sex ratios on different sized hosts. A model is constructed which includes the choice of accepting or rejecting a host as well as the choice of sex of offspring. The best reproductive strategy satisfies MacArthur's “product rule” for sex ratios and Charnov's “marginal value theorem” for optimal foraging. The model can be used to show that optimal sex ratio may vary with host density and size distribution.     

Manipulation of offspring sex ratio by second-mated female house wrens

In 1973, Trivers and Willard proposed that offspring sex ratio should be associated with the quality of parental care likely to be provided to the offspring. We tested this hypothesis by comparing fledgling sex ratios in nests of first- and second-mated female house wrens (Troglodytes aedon). In our Wyoming population, second-mated females typically receive little or no male parental assistance and fledge fewer and lower-quality young compared with first-mated females. Assuming that being of lower quality has stronger negative effects on the future reproductive success of males than that of females in this polygynous population, we predicted that fledgling sex ratios in the nests of second-mated females would be female-biased compared with the fledgling sex ratios of first-mated females. Additionally, we asked whether any sex bias at fledging could have resulted from male-biased nestling mortality caused by sex-biased parental provisioning. As predicted, mean fledgling sex ratios in nests of second-mated females were more female-biased than fledgling sex ratios in nests of first-mated females. However, we found no evidence of either sex-biased nestling mortality or sex-biased parental provisioning. These findings suggest that females are responding to their status as second-mated females and to the associated low-quality parental care that their young are likely to receive by producing female-biased clutches rather than manipulating the offspring sex ratio through sex-biased nestling mortality.     

Colony productivity and foundress behaviour of a native wasp versus an invasive social wasp

Abstract.  1. Colony productivity, prey utilisation, and foundress behaviour of a North American native wasp ( Polistes fuscatus ) versus an European invasive wasp ( Polistes dominulus ) were investigated in a controlled field experiment with optimal versus natural foraging conditions. Colonies with the optimal prey foraging conditions were provided with prey ad libitum within an enclosed area. The other colonies foraged in the adjacent field–woodland but had the same nest conditions as the other treatment.
2. When given prey ad libitum , both wasp species captured similar amounts of prey and the conversion to total offspring biomass was similar. But P. dominulus colonies produced 2.5 times the number of workers as P. fuscatus colonies, reflecting the smaller size of P. dominulus wasps.
3. Foundresses of P. dominulus were observed more often building or repairing the nest, thereby contributing to the production of colonies with twice as many cells as colonies of P. fuscatus . Foundresses of P. dominulus showed more acts of aggression toward workers than did P. fuscatus foundresses, which was not a function of adult density on the nest.
4. At the end of the experiment, P. dominulus colonies with optimal prey foraging conditions still had a high level of egg-laying and peaked in the number of pupae then, whereas egg-laying and the number of pupae per colony of the other treatments began to decline 2–3 weeks earlier. These results indicate that P. dominulus is more opportunistic than P. fuscatus , which may account in part for P. dominulus 's success as an introduced species in North America.     

Sublethal effect of brood parasitism on the grass-carrying wasp Isodontia mexicana

Abstract.  1. The nests of solitary, nest-provisioning wasps (Sphecidae and Crabronidae) are commonly attacked by brood parasites, including flies of the families Phoridae and Sarcophagidae. Larvae of the flies commonly kill the wasp offspring directly or starve it by consuming prey provided by the adult female wasps.
2. To determine whether brood parasitic flies can have a sublethal effect (i.e. reduced body size) on wasp offspring, nests of the wasp Isodontia mexicana were collected at two field sites in upstate New York, U.S.A. Nest diameter had no effect on the probability that a wasp cell would be inhabited by a brood parasite. Most offspring that developed in cells also containing phorids or sarcophagids managed to complete development and emerge as adults. Nevertheless, they had significantly smaller body size than conspecifics emerging from unparasitised cells in which the developing wasp did not have to compete for food (which in this species consists of tree crickets and katydids). Apparently, this is the first time that a sublethal effect of brood parasitism on offspring body size has been quantified for a solitary wasp species. Known effects of body size on the reproductive success of adult wasps suggest that sublethal consequences of the presence of brood parasites may have a significant effect on the fitness of adult I. mexicana .     

Red-winged blackbird predation on periodical cicadas (Cicadidae: Magicicada spp.): bird behavior and cicada responses

Summary Predation by red-winged blackbirds Agelaius phoeniceus L. on 13-year periodical cicadas (Magicicada spp.) and reactions by periodical cicadas to predators were studied during emergence of Brood XIX during summer of 1985 in northwestern Arkansas (USA). Emergences of periodical cicadas are classic examples of predator satiation due to high local densities of cicadas and birds are the major predators of adult periodical cicadas. Reactions of periodical cicadas to predators were assessed by recording behaviors exhibited by cicadas when approached in trees by a human hand during the 3-week period of peak adult densitics. Most male cicadas made a noise in association with escape behaviors when approached, and 50% of the females, which are silent, attempted escape behaviors. Observations of predation attempts on cicadas by red-winged blackbirds were made during the period of peak predation pressure. Red-winged blackbirds spent less time searching for cicadas over that 2-week period of increasing predation pressure, and became increasingly efficient at capturing cicadas. Handling time of cicadas by red-winged blackbirds increased by about 20 seconds over that period, as blackbirds spent more time consuming female cicadas. The flysquawk response, used only by male cicadas, was effective in deterring red-winged blackbirds; only 5% of the attacks by blackbirds were successful when that behavior was exhibited. All cicadas that remained motionless and silent when approached by blackbirds were captured and consumed. Because females remained inactive when approached more often than did males, blackbirds may have consumed more female cicadas. Changes that appeared in reactions of preiodical cicadas to the model predator and to the attacks of blackbirds reflect both changes in the sex ratio of the cicada populations and changes in behaviors of cicadas associated with mating and egg laying. The loud noise made by male periodical cicadas at mating centers did not appear to deter predation by blackbirds. Changes in the behavior of blackbirds that appeared to be in association with greater predation on female periodical cicadas relates directly to aspects of foraging theory, particularly predictions concerning more selective foraging during periods of abundant food resources.     

Does differential predation permit invasive and native mosquito larvae to coexist in Florida?

Abstract.  1. The hypothesis that selective predation on larvae of the invasive Aedes albopictus (Skuse) could account for its stable coexistence with the native mosquito species and inferior competitor Ochlerotatus triseriatus (Say) in Florida treeholes and container systems was tested experimentally.
2. Functional responses of the two dipteran predators Toxorhynchites rutilus (Coquillett) and Corethrella appendiculata (Grabham) were evaluated separately for A. albopictus and O. triseriatus prey. Both predators exhibited type II functional responses and consistently consumed more of the invasive species. Handling time of T. rutilus feeding upon O. triseriatus was significantly longer than when preying upon the invasive species.
3. When either predator species was offered varying ratios of the two prey species, A. albopictus was consumed preferentially. The absence of a prey ratio effect on preference indicated that switching probably does not occur.
4. The higher maximum feeding rate upon, and preference for, A. albopictus suggests that differential predation may foster coexistence of the invasive and native mosquito prey species in Florida.     

Chemical Mediation of Prey Recognition by Spider-Hunting Wasps

Predator–prey interactions are important in maintaining the structure and dynamics of ecological communities. Both predators and prey use cues from a range of sensory modalities to detect and assess one another; identification of these cues is necessary to understand how selection operates to shape predator–prey interactions. Mud-dauber wasps (Sphecidae) provision their larval nests with paralyzed spiders, and different genera of wasps specialize on particular spider taxa. Sceliphron caementarium (Drury 1773) wasps preferentially capture spiders that build two-dimensional (2D) webs, rather than those that construct three-dimensional (3D) webs, but the basis of this preference is not clear. Wasps may choose spiders based on an assessment of their web architecture, as 3D webs may provide better defenses against wasp predation than do 2D webs. However, because many hymenopterans use chemical cues to locate and recognize prey, it is also possible that mud-dauber wasps rely on chemical cues associated with the spider and/or the web to assess prey suitability. When we offered foraging S. caementarium wasps 2D and 3D spiders both on and off their webs, we found that in both cases the wasps took 2D spiders and avoided 3D spiders, demonstrating that the web itself is not the impediment. Results of a series of behavioral choice assays involving filter paper discs containing spider cues and chemically manipulated spiders or spider dummies corroborated the importance of spider chemical cues in mediation of prey recognition by mud-dauber wasps. We also discuss the relative importance of visual and chemical cues for prey recognition by wasps, examine the anti-predator behaviors of 2D and 3D spiders, and consider the role of wasp predation in spider diversification.     

Selective predation by larval Agabus (Coleoptera: Dytiscidae) on mosquitoes: support for conservation-based mosquito suppression in constructed wetlands

1. Wetland insect predators can structure aquatic prey communities via selective predation, but receive considerably less attention than vertebrate predators. We conducted laboratory experiments to test selective predation by two species of larval dytiscid beetles ( Agabus ; Coleoptera: Dytiscidae) and the potential contribution of these beetles to suppression of mosquito populations in constructed wetlands.
2.  Agabus consumed copepods, ostracods and mosquito larvae in no-choice tests. When offered a choice, 76% of all prey consumed were mosquito larvae, indicating selective predation. Subsequent experiments revealed this preference was due to ease of capture of mosquito larvae over alternative prey.
3. Cannibalism and intraguild predation were common within and between species of Agabus , which may reduce the overall impact of the observed selective predation.
4.  Agabus larvae selectively preyed on mosquito larvae over alternative prey, which is not characteristic of some fish used as biological control agents for mosquitoes. Predator exclusion or similar experiments in the field could document how these results translate into a natural setting.
5. The findings of this study suggest developing mosquito suppression strategies focused on conservation of native wetland predators. These strategies are preferable to introducing non-native generalist predators, or applying pesticides.     

Male-biased size dimorphism in ichneumonine wasps (Hymenoptera: Ichneumonidae) – the role of sexual selection for large male size

Abstract.  1. Sexual differences in body size are expected to evolve when selection on female and male sizes favours different optima.
2. Insects have typically female-biased size dimorphism that is usually explained by the strong fecundity advantage of larger size in females. However, numerous exceptions to this general pattern have led to the search for selective pressures favouring larger size in males.
3. In this study, the benefits of large size were investigated in males of four species of ichneumonine wasps, a species-rich group of parasitoids, many representatives of which exhibit male-biased size dimorphism.
4. Mating behaviour of all ichneumonine wasps are characterised by pre-copulatory struggles, in the course of which males attempt to override female reluctance to mate. A series of laboratory trials was conducted to study the determinants of male mating success.
5. A tendency was found for larger males as well as those in better condition to be more successful in achieving copulations. Size dimorphism of the species studied, mostly male-biased in hind tibia length but female-biased in body weight, indicates that sexual selection in males favours longer bodies and appendages rather than larger weight.
6. The qualitative similarity of the mating patterns suggests that sexual selection cannot completely explain the considerable among-species differences in sexual size dimorphism.
7. The present study cautions against using various size indices as equivalents for calculating sexual size dimorphism.
8. It is suggested that female reluctance in ichneumonine wasps functions as a mechanism of female mate assessment.     

Field observation of predation on paper wasp nests by introduced terrestrial slugs

Nest predation is an important factor in the colony mortality of paper wasps. Although a variety of animals are known to prey on paper wasp nests, colony failure of unknown cause often occurs in the field, and the existence of still-unknown predators is possible. Here, I report predation on nests of paper wasps, Polistes chinensis antennalis, by introduced terrestrial slugs, Lehmannia valentiana. The slugs were found on the wasps’ nests despite the presence of the owners on the nests and ate eggs, larvae, and presumably cell walls of the nests. This is the first report of predation on paper wasp nests by slugs.     

Interactions of the ant Crematogaster scutellaris with the fig/fig wasp mutualism

Abstract. 1. A classical example of specialised pollination mutualisms is the relationship between fig trees and their pollinating wasps, in which each partner depends completely on the other for its reproduction; however the fig/fig wasp association is also the target of a great diversity of other species, ranging from specialised parasites to opportunistic foragers, among them ants.
2. The ant community and the sources of ant attraction observed on the Mediterranean fig tree Ficus carica were characterised.
3. A guild of ants attracted by homopterans tended on the plant was distinguished from a second guild composed of two co-dominant ant species ( Crematogaster scutellaris and Pheidole pallidula ) that prey mostly on pollinating wasps, abundant during certain parts of the fig cycle.
4. Foraging workers of C. scutellaris search for prey on the fig inflorescence (syconium), capturing pollinating wasps mostly at the peak of wasp emergence and at a rate estimated to reach 600 prey per day for an entire tree.
5. Detailed study of the predatory sequences displayed under experimental conditions showed that ant workers captured 100% of the pollinating wasps offered, while they captured only 5.5% of the parasitoid wasp specific to the pollinator. The respective impacts of the interaction on ants and on the figs are discussed, as well as several behavioural traits of predation by the dominant ant on pollinators.     

The relevance of ants as seed rescuers of a primarily bird-dispersed tree in the Neotropical cerrado savanna

Sex ratios can influence mating behaviour, population dynamics and evolutionary trajectories; yet the causes of natural sex ratio variation are often uncertain. Although secondary (birth) sex ratios in guppies (Poecilia reticulata) are typically 1:1, we recorded female-biased tertiary (adult) sex ratios in about half of our 48 samples and male-biased sex ratios in none of them. This pattern implies that some populations experience male-biased mortality, perhaps owing to variation in predation or resource limitation. We assessed the effects of predation and/or inter-specific resource competition (intraguild predation) by measuring the local catch-per-unit-effort (CPUE) of species (Rivulus killifish and Macrobrachium prawns) that may differentially prey on male guppies. We assessed the effects of resource levels by measuring canopy openness and algal biomass (chlorophyll a concentration). We found that guppy sex ratios were increasingly female-biased with increasing CPUE of Macrobrachium, and perhaps also Rivulus, and with decreasing canopy openness. We also found an interaction between predators and resource levels in that the effect of canopy openness was greatest when Macrobrachium CPUE was highest. Our study thus also reveals the value of simultaneously testing multiple environmental factors that may drive tertiary sex ratio variation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Nest predators of woodland open-nesting songbirds in central Europe

I used time-lapse videotaping to identify predators of open songbird nests in fragmented deciduous woodland (nine plots, 2–10 ha each) in the Czech Republic from 2002 to 2006. I documented 22 species of predators at 171 nests of 13 species (mainly Blackcap Sylvia atricapilla , Song Thrush Turdus philomelos , Common Blackbird Turdus merula , Yellowhammer Emberiza citrinella and Chaffinch Fringilla coelebs ). The main predators were Pine Marten Martes martes (37% of 178 predation events), Jay Garrulus glandarius (29%), Buzzard Buteo buteo (7%) and Great Spotted Woodpecker Dendrocopos major (7%); mammals accounted for 48% of total predation. At least 3% of nests were depredated by multiple predators. In spite of their local abundance, Hooded Crows Corvus cornix did not present a serious threat for shrub nesting songbirds (< 1% of total predation). No predation by mice was recorded, suggesting that their importance has been overestimated in artificial nest studies. The proportional species composition of predators depended on which species occupied the monitored nest and location (study plot), but not on the year or the time of season. Corvids and raptors accounted for a relatively larger percentage of total predation of small ('warblers') and large ('thrushes') prey species, respectively, whereas carnivores were important predators of all prey species. Active nests of thrushes were only rarely robbed by Jays (< 4% of 52 events), presumably due to parental nest defence. Predation by woodpeckers was spatially clumped, probably due to individual foraging specialization. Predation by the other major predators was documented on most/all study plots.     

Identifying predators clarifies predictors of nest success in a temperate passerine

1.  Nest predation negatively affects most avian populations. Studies of nest predation usually group all nest failures when attempting to determine temporal and parental activities, habitat or landscape predictors of success. Often these studies find few significant predictors and interpret patterns as essentially random.
2.  Relatively little is known about the importance of individual predator species or groups on observed patterns of nest success, and how the ecology of these predators may influence patterns of success and failure.
3.  In 2006 and 2007, time-lapse, infrared video systems were deployed at nests of Swainson's warblers ( Limnothlypis swainsonii Audubon) in east-central Arkansas to identify dominant nest predators and determine whether factors predicting predation differed among these predators.
4.  Analysis of pooled data yielded few predictors of predation risk, whereas separate analyses for the three major predator groups revealed clear, but often conflicting, patterns.
5.  Predation by ratsnakes ( Elaphe obsoleta ) and raptors was more common during the nestling period, whereas predation by brown-headed cowbirds ( Molothrus ater ) occurred more during incubation. Additionally, the risk of predation by raptors and cowbirds decreased throughout the breeding season, whereas ratsnake predation risk increased.
6.  Contrary to expectations, predation by ratsnakes and cowbirds was more common far from edges, whereas raptor predation was more common close to agricultural edges.
7.  Collectively, our results suggest that associating specific predators with the nests they prey on is necessary to understand underlying mechanisms.     

Reconstruction of fig wasp mating structure: how many mothers share a fig?

Abstract.  1. Fig wasps (Hymenoptera: Agaonidae) represent an important model system for studies of sex ratio evolution, mainly because they may adjust their sex ratios in response to the numbers of ovipositing females (foundresses) that enter a fig and their clutch size.
2. Until recently, it was assumed that all foundresses fail to re-emerge from the figs that they have entered to oviposit, but there is increasing evidence that such re-emergence may be routine. The common practice of counting the number of dead foundresses present in a fig in order to deduce the number of foundresses is therefore questionable in species where failure to re-emerge has not been confirmed.
3. In this study, the alternative approach of microsatellite markers was used to reconstruct the within-fig breeding structure of a pollinating fig wasp by genetic analysis of the offspring. Broods of Liporrhopalum tentacularis , a species where foundresses regularly re-emerge from figs, were collected from figs of Ficus montana in their natural habitat in Indonesia as well as from an experimental glasshouse population in Leeds (U.K.).
4. The estimated foundress densities in the glasshouse population were similar to those in the field and ranged from one to six foundresses per brood.
5. Nearly 40% of all broods were produced by a single foundress, indicating that mating in these broods occurs exclusively between full siblings. High levels of inbreeding are therefore common in this species.     

Thermal and fitness-related consequences of nest location in Painted Turtles (Chrysemys picta)

1. Laboratory experiments have documented substantial temperature effects on the physiological ecology of reptilian eggs, embryos and offspring. However, functional links between important habitat characteristics, nest microenvironments and fitness-related traits of neonates in natural nests have rarely been studied.
2. A field study of 11 Painted Turtle ( Chrysemys picta ) nests was conducted to quantify the relationships between a habitat characteristic (i.e. vegetational cover around nests at oviposition) and (1) developmental temperature and its effect on offspring sex ratio and (2) hibernation temperature and its effect on offspring survivorship.
3. Vegetational cover was negatively correlated with nest temperatures in July, the period when offspring sex is determined. However, neither vegetational cover nor mean nest temperature predicted nest sex ratios, although correlations among these variables were consistent with causal relationships derived from laboratory studies.
4. Summer vegetational cover was also negatively correlated with measures of winter nest temperatures. Of the three nests exhibiting overwinter mortality, two were surrounded by thick vegetation and all experienced temperatures below – 8 °C. The remaining nests reached temperatures as low as – 6 °C without mortality, indicating that hatchlings in these nests exhibited remarkable supercooling ability.
5. The results suggest that habitat characteristics and nest microenvironments are functionally linked and have fitness consequences for both embryos and offspring, implying that nest-site choice by female turtles could have considerable utility.     

Life-history and sex allocation in Trypoxylon (syn. Trypargilum) lactitarse (Hymenoptera; Crabronidae)

Hymenopterans have become a model for the study of factors that govern sex allocation. In 1983, Seger proposed a model to study Sphecidae wasps with a strong prediction that sex ratio for univoltine wasps should be 1 : 1 (female : male), and for partially bivoltine species it should be male-biased. The present study investigates if Trypoxylon lactitarse (Saussure, 1867) is a univoltine or a bivoltine species and if Seger's model prediction fits the pattern of sex ratio found in this species. The study was carried out at Parque Municipal das Araucárias, in the municipality of Guarapuava, state of Paraná, southern Brazil, from December 2001 to December 2004. Nests of T. lactitarse were obtained using trap-nests drilled longitudinally to a depth to 80 mm with 7.0, 10.0 and 13.0 mm opening diameter. They were placed in a very heterogeneous site filled with araucaria forests, swamps and grasslands. Trypoxylon lactitarse showed two alternative life histories: either they pupated immediately and emerged as adults later in the same season (direct development), or they entered diapause, overwintering and pupating the following spring (delayed development). Its annual sex ratios were not significantly different from 1 : 1, and both sex ratio of overwintering and sex ratio of direct-developing wasps were also not significantly different from 1 : 1, in all years of this study. By examining these results, it was possible to conclude that although T. lactitarse is a multivoltine species, with four generations per year and two alternative life histories, its sex ratio did not support Seger's model.     

Host choice by Aphidius colemani: effects of plants, plant–aphid combinations and the presence of intra-guild predators

Abstract.  1. This study first measured the effect of plant (radish, pepper, and wheat), aphid–plant combination (the green peach aphid Myzus persicae on pepper and radish, and the bird cherry-oat aphid Rhopalosiphum padi on wheat) and the host on which Aphidius colemani was reared (the cotton aphid Aphis gossypii on cucumber, M. persicae on radish and pepper, and R. padi on wheat) on host choice behaviour of the parasitoid and the performance of its offspring. Then, the effect of predator presence ( Coccinella undecimpunctata larvae) on host preference of the wasps was tested.
2. When reared on M. persicae on either radish or pepper , wasps preferred the aphid–plant combination from which they had emerged. Wasps reared on A. gossypii (naïve to all hosts offered) and R. padi preferred to parasitise M. persicae on radish and M. persicae on either radish or pepper, respectively. Rhopalosiphum padi on wheat was the least preferred and also the least suitable host, as determined by offspring body size.
3. Contrary to expectations, the presence of predators did not influence the host choice of A. colemani , even when predator-free plants were offered nearby. Results indicate that wasps are more likely to remain in some host systems but not in others, even when facing risk of predation.
4. Results are discussed in respect to field data and the relative importance of risk of predation and host preference to wasp fitness.