Dominance and polyethism in the eusocial wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Dominance interactions affected patterns of non-reproductive division of labor (polyethism) in the eusocial wasp Mischocyttarus mastigophorus. Socially dominant individuals foraged for food (nectar and insect prey) at lower rates than subordinate individuals. In contrast, dominant wasps performed most of the foraging for the wood pulp used in nest construction. Social dominance also affected partitioning of materials collected by foragers when they returned to the nest. Wood pulp loads were never shared with nest mates, while food loads, especially insect prey, were often partitioned with other wasps. Dominant individuals on the nest were more likely to take food from arriving foragers than subordinate individuals. The role of dominance interactions in regulating polyethism has evolved in the eusocial paper wasps (Polistinae). Both specialization by foragers and task partitioning have increased from basal genera (independent-founding wasps, including Mischo-cyttarus spp.) to more derived genera (swarm-founding Epiponini). Dominance interactions do not regulate forager specialization or task partitioning in epiponines. I hypothesize that these changes in polyethism were enabled by the evolution of increased colony size in the Epiponini. Received: 8 December 1997 / Accepted after revision: 28 March 1998     

Parental investment and its ecological consequences in the solitary wasp Sceliphron assimile (Dahlbom) (Sphecidae)

Summary The sphecid wasp Sceliphron assimile Dahlbom builds mud cells whose volumes are positively correlated to maternal length, the relationship being highly significant for daughters (Fig. 1). However, small females build cells longer and wider in relation to their own body length than large females do (Fig. 3). Cell volume closely regulates the total weight of prey stocked per cell (Figs. 4, 10). Maternal length is directly related to prey size and inversely related to the number of prey stocked (Fig. 5). Female length, above a threshold of 19 mm, probably does not influence her chance of starting a nest (Fig. 6) but is strongly positively correlated to achieved fecundity. Large females have larger offspring and a higher proportion of daughters than small females do (Fig. 7), but this is largely controlled by maternal behaviour rather than by genetical mechanisms (Fig. 10). The results are in general agreement with those of Trivers and Hare (1976) for solitary wasps, but large progeny are cheaper to produce per unit weight than small ones, and for a given weight sons are cheaper to produce than daughters.     

Sex allocation in the solitary bee Osmia cornuta: do females behave in agreement with Fisher's theory?

Fisher's theory of sex allocation predicts that, in a panmictic population, parental investment will be equally distributed between male and female progeny. Most studies on parental investment in nesting solitary bees and wasps use offspring or provision weight as estimators of parental investment and do not corroborate Fisher's theory. The measurement of parental investment may be confounded by several factors. First, the use of offspring or provision size does not account for seasonal variation in foraging costs associated with aging of nesting females. Second, provision or offspring size do not reflect parental investment associated with nest construction. In this two-year study we measured parental investment in a solitary bee. We calculated sex allocation using both provision weight and foraging time as parental investment estimators. Investment in pollen-nectar provisions decreased, while investment in mud structures (nest construction) increased, as the nesting period progressed. Overall investment in provisions per nest was ∼25 times higher than investment in mud. Pollen-nectar foraging trips became longer as the season progressed, but mud trip duration did not vary. Due to weather differences between years, more offspring per female were produced in the first year, but progeny sex ratio and mean offspring size of both sexes were similar between years. Mortality did not differ between sexes. As predicted by Fisher's theory, production cost ratios did not differ from 1 in either year, irrespective of the currency used to estimate parental investment (provision weight or foraging time). Our results strongly support Fisher's theory.     

Effects of prey quantity on predatory wasps (<Emphasis Type="Italic">Polistes dominulus</Emphasis>) when patch quality differs

To determine the effects of prey quantity on central-place foraging of predatory wasps ( Polistes dominulus), prey of varying quality were distributed in patches. A field experiment was conducted, which controlled the amount and quality of prey available. 'Low-fed' colonies were provided with one-third the quantity of prey as that of 'high-fed' colonies. Both were provided with a 1:1 ratio of palatable:unpalatable prey. Experienced wasps of the high-fed colonies never selected unpalatable prey as their first choice of the day, whereas those of the low-fed colonies selected unpalatable prey as their first choice about 1:4 times. In general, wasps from the high-fed colonies reduced palatable patches to zero prey before exploiting unpalatable patches. Foundresses of high-fed colonies captured disproportionately more palatable prey than those of low-fed nests, but there was no correlation between ratio of palatable to unpalatable prey taken and the number of offspring produced. Wasps from low-fed colonies attacked unpalatable prey sooner, but not without considerable effort to avoid use of those patches. Foundresses of low-fed colonies also spent a greater proportion of time overall in unsuccessful search, which may explain why only wasps from low-fed colonies foraged on cool days. High-fed colonies produced more cells and more and heavier offspring than low-fed colonies. But the productivity of the low-fed colonies was greater than that of the 'natural' colonies, which had to find their own prey in a field-woodland area. These results indicate that prey scarcity changes foraging behavior and affects prey choice. These changes may not totally alleviate negative effects of unpalatable prey on colony development and offspring production. The results of this study increase understanding of the central-place foraging behavior of paper wasps, which are important biocontrol agents in natural and agricultural settings.     

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     

Relationship between body size, provisioning rate, longevity and reproductive success in females of the solitary bee Osmia cornuta

Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.     

Parental investment in two solitary wasps Ancistrocerus adiabatus and Euodynerus foraminatus (Eumenidae: Hymenoptera)

Summary Females of the solitary eumenid wasps Ancistrocerus adiabatus and Euodynerus foraminatus control the adult size of their offspring by the amount of food provded to the larvae. For both species, larger females provision more offspring and collect more food than smaller females. Males of E. foraminatus, upon emergence as adults, fight for control of the nest area. The winning brother remains at the nest and mates with his sisters as they emerge several days later. Males of A. adiabatus are not as agonistic as males of E. foraminatus so that a number of males may remain near the natal nest. When females nest near each other, male-male interactions increase, and the likelihood of sibling mating decreases. Under conditions of clumped nesting, females make larger offspring and invest more in males than when nesting in isolation.     

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.     

Brood-provisioning strategies in Dawson's burrowing bee, Amegilla dawsoni (Hymenoptera: Anthophorini)

Males of Dawson's burrowing bee are dimorphic in size. Although large (major) males defeat smaller ones in competition for emerging females and therefore are more likely to mate, majors are greatly outnumbered by half-sized (minor) males. Nesting females might produce many minor males, despite their low reproductive value, because female behaviour is governed by a mixed evolutionarily stable strategy (ESS), in which case the ratio of majors to minors should not be affected by changes in female condition. In contrast, a conditional-strategy hypothesis predicts that older, wing-worn or stressed females unable to forage efficiently should be especially likely to produce minor offspring, which require less brood provisions. To test these alternative hypotheses, we manipulated the condition of nesting female bees by the addition of weights and the removal of their wing margins. These manipulations, done early in the flight season, failed to increase the production of minor males, a result consistent with the mixed-ESS hypothesis. However, unmanipulated females were far more likely to produce minor males if they were small or if they were nesting late in the season, when foraging conditions had deteriorated, findings that are consistent with a conditional provisioning strategy. Thus it appears that the abundance of minor males is the result of a conditional provisioning strategy of nesting females, which may be superimposed on a fixed tendency to produce large offspring early in the season and small ones later.     

Compensating for the costs of polygyny in hen harriers Circus cyaneus

In polygynous species, the adults are faced with a dilemma during chick rearing. Males must decide how to distribute food between their females and food allocation patterns are often highly unequal. In turn, the females that receive less food from males have to decide how much time to invest in additional hunting. If they spend more time hunting, then they leave their young exposed to weather and predators. However, if they stay at the nest, they increase the risk of their chicks starving. One way that birds may compensate for reduced provisioning is by increasing the size of prey caught. We tested this hypothesis by comparing prey deliveries to nests of hen harriers, Circus cyaneus, with females of different breeding status. As expected, male harriers delivered less food items to the nests of polygynous females, and especially their secondary, or β females. However, both sexes were able to compensate by delivering larger items and there was no difference in the overall mass of food delivered to nests. Moreover, females spent a similar amount of time at the nest, irrespective of status, and there were no overall differences in breeding success. Our results show that polygynous female harriers can compensate for the costs of polygyny, but we suggest that their ability to do so will vary according to the abundance of both large prey and predators.     

Host choice and offspring sex allocation in a solitary parasitic wasp

Summary Females of the parasitic wasp Antrocephalus pandens can detect differences in the quality of their hosts (pupae of Corcyra cephalonica, a stored-product moth) and allocate offspring of either sex accordingly. Larger and younger hosts are accepted more often in both dead and live hosts; more female offspring emerge from the perceived better hosts, while more males emerge from the smaller, older ones. These patterns are consistent with a sex allocation strategy by the mother, since females from a given size host tend to be larger than males and larger females produce more eggs. However, when wasps lay their eggs in groups of hosts of different size and age rather than encountering them one at a time, no difference in number or sex ratio of offspring is detected between groups. This result and evidence from the change in offspring sex ratio with female age and with numbers of females foraging on a group of hosts are interpreted and discussed in the context of sex allocation (Charnov 1979) and local mate competition (LMC, Hamilton 1967) theories.     

Mate quality,not aggressive spillover,explains sexual cannibalism in a size-dimorphic spider

Sexual cannibalism particularly before mating is costly for the male victim but also for the female aggressor if she risks remaining unmated. The aggressive spillover hypothesis explains the persistence of this behavior as a maladaptive side effect of positive selection on aggressiveness in a foraging context. The hypothesis predicts that the occurrence of sexual cannibalism is explained by female aggressiveness but is not related to male phenotype or behavioral type. An alternative hypothesis invokes sexual selection and makes the opposite prediction namely that sexual cannibalism is an expression of female choice and should hence mainly target males of low quality. We tested the above hypotheses on a sexually dimorphic nephilid spider Nephilengys livida, known for male monopolization of females via genital damage, female genital plugging, and mate guarding, by staging mating trials during which we recorded mating behaviors and occurrences of pre- and postcopulatory cannibalism. We did not restrict assessment of aggressiveness to the mating and foraging context but also included aggression against same sex conspecifics. To assess female personalities, i.e., consistent individual differences in behavior including aggressiveness, we repeatedly tested them for intra-sex aggression, voracity towards prey, locomotory activity, and boldness. Females exhibited consistent differences in intra-sex aggressiveness, latency to attack prey, and boldness. Aggressive females had shorter latencies to attack prey and were more active than non-aggressive ones. In contrast to the predictions of the aggressive spillover hypothesis, females that were aggressive towards prey and towards other females were not more likely to attack a male than non-aggressive females. In support of the mate choice hypothesis, less aggressive males were more likely attacked and cannibalized than more aggressive ones. This hints at sexual selection for aggressiveness in males and raises the question of mechanisms that maintain variation in male aggressiveness.     

Inter-annual changes in diet and foraging trip lengths in a small pelagic seabird, the thin-billed prion Pachyptila belcheri

Central place foragers are constrained in their foraging distribution by the necessity to return to their nest site at regular intervals. In many petrels that feed on patchily distributed prey from the sea surface over large foraging areas, alternating long and short foraging trips are used to balance the demands of the chick with the requirements of maintaining adult body condition. When the local conditions are favourable for prey density and quality, adults should be able to reduce the number of long foraging trips. We studied the flexibility in foraging trip lengths of a small pelagic petrel, the thin-billed prion Pachyptila belcheri, over three breeding seasons with increasingly favourable, cold-water conditions. During a warm-water influx in February 2006, chicks were fed less frequently and adults carried out foraging trips of up to 8 days. When conditions became more favourable with colder water temperatures in 2007 and 2008, thin-billed prions decreased trip lengths, more often attended their chick every day, and long foraging trips of six to eight days were not registered during 2008. Chick growth rates mirrored this, as chicks grew poorly during 2006, intermediate during 2007 and best during 2008. Thin-billed prions preyed mainly on squid during incubation and mainly on amphipods and euphausiids during chick-rearing. In the poorest season only, the diet was substantially supplemented with very small copepods. Together, the present results indicate that during warm-water conditions, thin-billed prions had difficulties in finding sufficient squid, amphipods or euphausiids and were forced to switch to lower trophic level prey, which they had to search for over large ocean areas.     

The effect of maternal body size on embryo survivorship in the broods of pregnant male pipefish

The occurrence of male pregnancy in the family Syngnathidae (seahorses, pipefishes, and sea dragons) provides an exceptionally fertile system in which to investigate issues related to the evolution of parental care. Here, we take advantage of this unique reproductive system to study the influence of maternal body size on embryo survivorship in the brood pouches of pregnant males of the broad-nosed pipefish, Syngnathus typhle. Males were mated with either two large females, two small females, a large then a small female, or a small then a large female. Our results show that offspring survivorship depends on an interaction between female body size and the number of eggs transferred by the female. Eggs of larger females deposited in large numbers are more likely to result in viable offspring than eggs of smaller females laid in large numbers. However, when females deposited smaller numbers of eggs, the eggs from smaller females were more likely to produce viable offspring compared to those from larger females. We found no evidence that this result was based on mating order, the relative sizes of competing females, or egg characteristics such as dry weight of eggs. Additionally, male body size did not significantly influence the survivorship of offspring during brooding. Our results suggest that the factors underlying offspring survivorship in pipefish may be more complex than previously believed, with multiple factors interacting to determine the fitness of individual offspring within the broods of pregnant males.     

Different habitats, different habits? Response to foraging information in the parasitic wasp Venturia canescens

The parasitic wasp, Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), has two reproductive modes, namely, thelytoky or arrhenotoky, and occurs in habitats with highly variable or relatively stable host abundances, respectively. Since information processing is costly, we expected that information indicating resource availability would be mainly used by the thelytokous wasps and less so by the arrhenotokous type. This idea was explored by two different approaches. In a study on patch-time allocation, we used females from ten populations and measured patch-residence times of individuals that visited multiple patches at different encounter rates. In a more detailed approach, thelytokous and arrhenotokous females from a single location were observed continuously while foraging, and all behaviors were recorded. Wasps of both reproductive modes (i.e., both habitat types) used information for the assessment of habitat quality. However, the way that the information was used differed between them. Whereas thelytokous females used foraging information to maximize their efficiency at high patch-encounter rates, arrhenotokous females merely reduced the number of offspring produced without changing patch times. The behavior of the arrhenotokous females should result in a spreading of offspring across the habitat and, thus, reduced sib-mating. The foraging strategy of these wasps might therefore be an adaptation to reduce costs associated with inbreeding.     

Foraging costs of a tail ornament: Experimental evidence from two populations of barn swallows Hirundo rustica with different degrees of sexual size dimorphism

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.Communicated by M. Zuk     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

The influence of prey distribution on the foraging strategy of the lizard Oligosoma grande (Reptilia: Scincidae)

The grand skink, Oligosoma grande, is a diurnal rock-dwelling lizard from the tussock grasslands of Central Otago, New Zealand, whose diet includes a variety of arthropods and fruit. We conducted a field experiment to examine the influence of prey distribution on foraging behavior and spacing patterns. On sites where prey distribution was unaltered (control sites), males and females differed in diet and foraging behavior. Most male feeding attempts were directed at large strong-flying insects, and males used a saltatory search pattern that involved relatively infrequent moves of long duration. Females spent more effort catching small weak-flying insects and visiting fruiting plants. Their search behavior involved frequent moves of short duration. The placement of meat-bait on experimental sites led to a redistribution of large flies without influencing other prey types. Experimental females switched foraging strategy by adopting a search pattern of relatively infrequent moves of long duration, increasing the frequency of attempts to capture large prey, and reducing the importance of fruit in their diet. The experimental manipulation appeared to influence space use. On control sites, both sexes had comparably sized home ranges. On experimental sites, male home ranges were significantly larger than female home ranges. Received: 3 November 1997 / Accepted after revision: 13 December 1998     

Predatory behavior of Polybia sericea (Olivier), a tropical social wasp (Hymenoptera: Vespidae)

Summary Polybia sericea (Olivier) (Hymenoptera: Vespidae) foragers were trained to visit experimental foraging plots and tests were conducted to determine the role of visual, olfactory, and chemotactile cues in prey location. Foragers approached prey from downwind and hovered downwind of visual and olfactory stimuli. Olfactory cues were more likely to elicit landing than were visual cues. Biting of potential prey was most consistently elicited by a combination of visual, tactile, and chemotactile cues. Foragers encountering large prey carried a piece back to the nest and returned for prey remains. These returning foragers used visual cues to direct intensive aerial search; olfactory prey cues elicited landing.     

The organization of work in Polybia occidentalis: costs and benefits of specialization in a social wasp

Summary Nest construction, a complex social activity requiring the coordination of 3 tasks (Fig.2), was compared in large (<350 adults) and small (<50 adults) colonies of Polybia occidentalis. The 3 tasks—water foraging, pulp foraging, and building—are performed by 3 separate groups of workers (Fig.4). Of the 8 acts comprising the 3 tasks, 5 regularly involve the transfer of water or pulp from one worker to another on the nest.Small colonies required nearly twice as long (35.4 worker-min) as large colonies (20.1 workermin) to complete a unit amount of construction work. Behavioral acts involving material transfer among workers were responsible for most of the increase in small colonies. In other words, the waiting times experienced by material donors and recipients were greater in small colonies. In small colonies workers switched among the three tasks more frequently than in large colonies (Fig. 4). This was the result of more frequent switching by generalists (workers that performed 2 or 3 of the tasks), rather than by a decrease in the proportion of specialists (workers performing only 1 task type) (Fig. 3).The series-parallel system by which Polybia occidentalis organizes nest construction has a major advantage over the series operation of solitary wasps. Pulp foragers collect and carry loads that are 6.1 times as large as builders can work with at the nest, and water foragers bring in loads that appear to be limited only by crop capacity and that provide all the moisture necessary for the complete processing of 0.74 of a foraged pulp load. As a result P. occidentalis can collect and process a given amount of nest material using 2.6 times fewer foraging trips than would be required by the series system. This in turn means that P. occidentalis not only achieves an energy saving that probably more than offsets the increased costs of material handling at the nest, but it reduces the exposure of its foragers to predators in the field.