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

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

Individual variation and prior experience affect the discrimination of a brood-parasite by its subsocial beetle host

In birds, host experience can modify response to parasites but nothing similar is known for insects. We studied two desert tenebrionid beetles, the subsocial host Parastizopus armaticeps and its obligate and ubiquitous brood-(clepto)parasite Eremostibes opacus, which mimics the hosts odour spectrum. Nearly 10% of host burrows in the field (n=214) remain unparasitised even after introducing E. opacus experimentally (n=22). In discrimination tests, 7% of naive host beetles eject the parasite, showing individual variation in olfactory discrimination ability. To test for effects of prior breeding experience, naive (first time) or experienced (second time) breeders with or without prior parasite contact were confined with E. opacus at breeding onset, their behaviour recorded and parasite number counted on breeding completion. More first- than second-time breeding pairs completely excluded the parasite and second-time breeders with no prior contact with E. opacus had fewer parasites than any others. These pairs increased burrow-guarding dramatically, mostly during early reproduction, this being the best predictor of parasite number. Prolonged guarding, however, must be traded-off against other brood-care behaviours. Hence, it might pay breeding pairs to take the risk of accepting some parasites breeding within their burrow, and we expect this effect will be more pronounced if poor feeding and digging conditions are taken into account. This might explain why apparently 90% of the P. armaticeps breeding burrows in the field contain at least one parasitic E. opacus.Communicated by D. Gwynne     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Pair and extra-pair mating success relative to male quality in red-winged blackbirds

We tested whether the reproductive success of male red-winged blackbirds (Agelaius phoeniceus) varied with male secondary sexual traits or with haematozoa prevalence, and whether these patterns were consistent with females preferring genetically superior males. We also determined whether the traits that correlated with male success on their own territories were also correlated with male success at siring young on other males territories. Our analysis included data from a 6-year study involving 617 nestlings for which paternity was determined by DNA profiling. Larger males sired more young on their territories, principally because they obtained larger harems. The success of larger males at acquiring more mates did not appear to be a consequence of larger males holding larger or better-quality territories. Older and longer-lived males sired more young by extra-pair fertilizations. Larger males sired the most offspring overall (on territory + off territory). Variation in epaulet size and color, responses to male and female models, nest defence and parasitism was not correlated with male success either on or off their territories or overall. Male success in a given year was significantly correlated with success the previous year, as expected if females were selecting genetically superior males. The male that was by far the most successful individual in this study was highly consistent from year to year. Because male body size is positively correlated with survival in this population (although not within the sample of males included in this study), female preference for larger males may have reflected a general preference for males with superior Survival ability. We propose that the direct advantage realized by older males in extra-pair matings might indicate that experience is important, such that experienced males are better at creating or exploiting the opportunities for extra-pair mating. This hypothesis is consistent with a pairwise analysis of cuckoldry that showed that cuckolders were most often older than the males they cuckolded.     

Nestling size rank in the little egret (Egretta garzetta) influences subsequent breeding success of offspring

Few studies have investigated the long-term fitness consequences of nestling size hierarchies in altricial birds. In this study, we investigated whether or not the size rank order of siblings influences subsequent breeding success in the little egret, Egretta garzetta. From a marking program allowing individual recognition of wild birds, we obtained data on the breeding success of 56 pairs comprising individuals for which the size rank order was known. The breeding success in these pairs was positively influenced by the age of the marked bird but negatively affected by the laying date of the pair and the size rank order of the marked individual. There was also a significant difference between breeding colonies. We suggest two main hypotheses for a link between size rank order of individuals and their breeding success and we discuss our results in relation to current hypotheses on the adaptive value of hatching asynchrony. Received: 10 August 1998 / Accepted after revision: 13 December 1998     

Cooperative breeding increases reproductive success in the social spider <Emphasis Type="Italic">Stegodyphus dumicola</Emphasis> (Araneae,Eresidae)

Sociality in some birds, mammals, and social insects was suggested to have evolved through the lengthening and extension of parental care behaviors to nondirect descendents. In these systems, group members care for young cooperatively and, thus, increase the reproductive success of the breeders and fitness of the young. Parental care behaviors, such as regurgitation feeding and matriphagy (consumption of the mother), occur in several subsocial and social spiders. However, it is not known whether females in a colony cooperate in caring for the young of other females and whether such cooperative care improves reproductive success. To answer this question, we created experimental colonies of the social spider Stegodyphus dumicola (Araneae, Eresidae), allowing only one female in a group to produce young, simulating reproductive skew occurring in nests in nature. In this paper, we show for the first time that females of S. dumicola cooperate in providing regurgitated food for young of other females and are even eaten by those young. Young raised by a group of females were larger and had greater survival than young raised only by their mother. Thus, fitness benefits from raising broods cooperatively may have favored the evolution of sociality in spiders.     

Modified pheromone communication associated with insecticidal resistance in the obliquebanded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

Summary. This study reports on the impact of insecticidal resistance on the diel periodicity of the calling behaviour and pheromone production of different-aged virgin females of the obliquebanded leafroller (OBL), Choristoneura rosaceana. While both resistant (R) and susceptible (S) females initiated calling on the first night following emergence, the periodicity of the calling behaviour, as determined by the mean onset time of calling (MOTC) and the mean time spent calling (MTSC) over the first six nights of calling, differed between the two strains. R females started calling significantly later in the night. However, as the MOTC of R females advanced with age but did not do so in S individuals, the difference between strains was more pronounced in younger than older females. Furthermore, R females spent less time calling than S individuals. However, the MTSC increased as a function of age in both R and S females, so the difference between strains remained fairly constant for each night of calling. The major component of OBL sex pheromone, the Z11-14: Ac, determined at peak calling activity, significantly declined with female age. Overall, pheromone production was lower in R females than in S females, with the difference being more pronounced in younger than in older individuals. Thus, resistant females may have a lower mating success. The mating success of both R and S strain males did not vary with the number of previous matings acquired. With regard to males, although there was a significant decline in spermatophore size with successive matings, there was no significant difference between strains. However, R males are smaller and may be disadvantaged through female choice and/or may respond differently to pheromone source compared with S individuals. If the reproductive success of both sexes is affected, this may have a profound influence on the dynamics of insecticidal resistance in the presence or absence of selection in OBL populations. Received 4 July 2001; accepted 19 October 2001.     

Sex-specific differences in site fidelity and the cost of dispersal in yellow-headed blackbirds

Male migratory birds tend to be more faithful than females to previous breeding sites, suggesting sex differences in costs or benefits of dispersal. In Illinois, greater site fidelity by male yellow-headed blackbirds (Xanthocephalus xanthocephalus) was associated with reduced reproductive success the following year for dispersers relative to non-dispersers. Dispersing females suffered no reduction in reproductive success the following year. Males that attracted few social mates, and thus had low reproductive success, were more likely to disperse, whereas females dispersed in response to low-patch reproductive success, regardless of their individual performance. Males that dispersed appeared to be successful acquiring territories because none was observed as a floater. The rate of dispersal by males in this low-density population was greater than in more dense populations where dispersing males may be less successful at acquiring territories. Despite success at obtaining territories, males that dispersed acquired territories on the periphery of wetlands where fewer females nested, resulting in lower reproductive success. In the second year after dispersing, however, males moved onto more central territories where they acquired larger harems. Thus, dispersal by males may be a long-term strategy requiring at least 2 years for benefits to be realized. Long-term success was enhanced because dispersing males moved to wetlands on which reproductive success was higher than on the wetlands they left. In addition to demonstrating that both individual and patch reproductive success affect dispersal decisions, these data indicate that when evaluating costs and benefits of dispersal, researchers should use a time frame beyond 1 year.     

Philopatry and mother-daughter associations in bushy-tailed woodrats: space use and reproductive success

Social groups in mammals are often based on overlapping generations of philopatric females, yet few researchers have examined fitness consequences of associations between females and their daughters. We examined survival and reproductive consequences of sharing a natal site with one's mother or daughter in wild bushy-tailed woodrats (Neotoma cinerea). Bushy-tailed woodrats reside on discrete rocky outcrops, and females tend to spend their entire reproductive life on their natal outcrop. Mothers and daughters that co-existed on the natal outcrop were closely associated in space. Juvenile females were more likely to survive their first winter to become breeding yearlings at their natal site if their mother was also present at the natal site during the winter. Juveniles survived equally well with or without their mother where densities of adult females were experimentally reduced, suggesting that females enhance their daughter's survival by facilitating their access to limited resources. Yearling females exhibited no reproductive costs from association with their mothers at the natal site during the breeding season; in fact, yearlings that bred at the natal site simultaneously with their adult mothers tended (P = 0.07) to experience fewer reproductive failures than did those that bred in the absence of their mothers. Mothers did not experience any detectable costs when sharing a natal site with a daughter. We conclude that bushy-tailed woodrats exhibit stable, fitness-enhancing associations among mothers and their philopatric daughters.