Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.     

Risk taking during parental care: a test of three hypotheses applied to the pied flycatcher

According to life-history theory, there will often be a conflict between investment in current versus future reproduction. If a predator appears during breeding, parents must make a compromise between ensuring the growth and survival of offspring (nest defence, feeding and brooding of young), and reducing the risk of predation to ensure their own survival. We model three hypotheses for the outcome of this conflict which are particularly relevant for altricial birds. They are not mutually exclusive, but focus on different costs and benefits. (1) Parental investment is determined by the parents’ own risk of predation. This hypothesis predicts that a lone parent should take smaller risks than a parent that has a mate. (2) Parental investment is related to the reproductive value of the offspring: Parents are predicted to take greater risks for larger broods, larger-sized or older offspring. (3) Finally, we present the new hypothesis that parental investment is related to the harm that offspring would suffer during a period of no parental care (incubation, brooding, feeding). This hypothesis predicts that parents should take greater risks for younger offspring, or for offspring in poorer condition, because the marginal benefit of parental care is largest in such cases. Hence, one may also expect that lone parents should take greater risks than two parents because their offspring are more in need of care. We tested these hypotheses on the pied flycatcher (Ficedula hypoleuca) by presenting a stuffed predator of the parents (a sparrowhawk, Accipiter nisus) close to the nest when parents were feeding the young. Risk taking was measured as the time that elapsed until the first visit to the nest. Most support was found for the ‘‘harm to offspring’’ hypothesis. Previous studies have usually measured the intensity of nest defence against typical nest predators, and have found evidence for the ‘‘reproductive value of offspring’’ hypothesis. However, our model predicts that the importance of the reproductive value of the offspring should decrease relative to the harm that offspring would suffer if they were not cared for when the predator type changes from a nest predator to a predator of adults, and when conditions for breeding turn from good to bad. Received: 13 April 1995/Accepted after revision: 11 March 1996     

Preparation and activation: determinants of age at reproductive maturity in male baboons

Age at maturity is a particularly important life history trait, but maturational data are rare for males in natural populations of mammals. Here we provide information on three maturational milestones and their social and demographic correlates among 43 wild male baboons, Papio cynocephalus, in a natural population in Amboseli National Park, Kenya. We examined (1) age at testicular enlargement, which signals puberty and the onset of subadulthood, (2) age at attainment of adult dominance rank, which we consider to be the beginning of adulthood, and (3) age at first sexual consortship, which is the best measure available for age at first reproduction in male baboons. Testicular enlargement (median age = 5.69 years) occurred earlier among sons of high ranking mothers, and was not influenced by rainfall or seasonality. Attainment of adult dominance rank (median age = 7.41 years) was also accelerated among sons of high-ranking mothers, and among males whose mothers had died while the males were juveniles. First sexual consortship (median age = 7.92 years) was not influenced directly by maternal characteristics, but attainment of adult dominance rank always preceded first consortship. The lag time between attainment of adult rank and first consortship (median = 2.5 months; range = 5–526 days), was predicted by the number of sexually cycling females in the group when the male attained rank, and by how high ranking the male became in his first months as an adult. We suggest that the age at which a male baboon is ready to begin reproducing is influenced by a relatively stable maternal characteristic that exerts its influence early in development, but the timing with which this potential is realized depends on activation by more proximate, often stochastic triggers such as female availability. This two-level organization of influences is likely to contribute to the variance both in age at first reproduction and in lifetime fitness. Differences in the relative magnitude of the two levels will lead to both intra- and interspecific variability in the opportunity for maternal selection and sexual selection.     

Reproductive effort of colonial and solitary breeding tree sparrows Passer montanus L.

A total of 250 nestboxes were arranged in five plots in a suburban area of Budapest, Hungary (19°04E, 47°41N). In each plot, 25 were placed at 50 m intervals to simulate solitary breeding and 25 3–5 m apart to simulate colonial breeding. Length of nest building period, feeding frequency, nestling mortality, nestlings' diet, productivity and parental condition were compared for colonial and solitary breeding tree sparrows Passer montanus. Parents with long nest-building periods, including the majority of first-year females, produced fewer young than parents which built over short periods. Parents fed nestlings morefrequently and nestlings had lower mortality in second than first broods; whether or not a third brood was reared was determined by the costs invested in first and second broods. Females that laid a third clutch had reared fewer young in first and second broods and were heavier than females that reared many young in two broods. Colonial birds had higher feeding frequencies, more similar diets and suffered lower nestling mortality than solitary parents for first broods, but they fed less frequently, diets were less similar, and nestling mortality was higher in second and third broods. It is suggested that colonial breeders benefited from the social stimulation of simultaneous feeding in first broods, but the advantage of synchronicity in feeding declined in second broods and the sparser breeding spacing of solitary parents was more advantageous for feeding in second and third broods. Birds that changed nest spacing between broods had fed nestlings less frequently and had higher nestling mortality before changing than birds which retained their spacing. Parents which changed from colonies to solitary nests fed more frequently with lower nestling mortality in the next brood than parents which retained colonial nests for their second (and third) brood. Solitary parents did not show such a relationship. The rearing of three broods caused higher weight loss in colonial than solitary parents.Correspondence to: L. Sasvári     

Delayed dispersal as a potential route to cooperative breeding in ambrosia beetles

Xyleborini are a species-rich tribe of ambrosia beetles, which are haplodiploid and typically mate among siblings within their natal brood chamber. Several characteristics of this tribe would predict the evolution of higher levels of sociality: high genetic relatedness within galleries due to inbreeding, high costs of dispersal and the potential benefit of cooperation in brood care within the natal gallery (e.g. by fungus gardening, gallery extension, offspring feeding and cleaning). However, information on the social system of these beetles is very limited. We examined the potential for cooperative breeding in Xyleborinus saxeseni by monitoring dispersal in relation to brood size and composition. Results show that adult female offspring delay dispersal despite dispersal opportunities, and apparently some females never disperse. The females’ decision to stay seems to depend on the presence of eggs and dependent siblings. We found no indication that female offspring reproduce in their natal gallery, as colonies with many mature daughters do not contain more eggs than those with few or no daughters. There is a significant positive relationship between the number of females present and the number of dependent siblings (but not eggs), which suggests that cooperative brood care of female offspring raises colony productivity by improving survival rates of immatures. Our results suggest that cooperative breeding is likely to occur in X. saxeseni and possibly other xyleborine species. We argue that a closer look at sociality within this tribe may yield important information on the factors determining the evolution of cooperative breeding and advanced social organization.     

Female benefit,male risk: Polyandry in the true armyworm Pseudaletia unipuncta

In Lepidoptera polyandry is common and females may increase their lifetime reproductive output through repeated matings if they acquire essential resources from male ejaculates. However, the paternity of males mating with previously-mated females is far from assured unless sperm precedence is absolute. In this study on the polyandrous armyworm, Pseudaletia unipuncta, we used two strains of male (the black-eyed wild type and a red-eyed homozygous, recessive mutant), mated with red-eyed females, to determine (i) whether male investment has any impact on female reproductive output, and (ii) if females do benefit from multiple matings, to what extent males fertilize the eggs to which they contributed. Multiple mating resulted in a significant increase in both the fecundity and longevity of females. However, the degree of sperm precedence (those eggs fertilized by the second male) varied from 0–100%, but was not affected by either male size or age, or by the duration of copulation. In cases where sperm precedence was <50% (x = 12%) females produced significantly more eggs (1384 versus 940) prior to the second mating than females where sperm precedence was >50% (x = 89%), indicating that the quality of the first mating influenced the fertilization success of the female's second mate.     

A test of reproductive skew models in a field population of a multiple-queen ant

Determining the evolutionary basis of variation in reproductive skew (degree of sharing of reproduction among coexisting individuals) is an important task both because skew varies widely across social taxa and because testing models of skew evolution permits tests of kin selection theory. Using parentage analyses based on microsatellite markers, we measured skew among female eggs (n=32.3 eggs per colony, range=20–68) in 17 polygynous colonies from a UK field population of the ant Leptothorax acervorum. We used skew among eggs as our principal measure of skew because of the high degree of queen turnover in the study population. Queens within colonies did not make significantly unequal contributions to queen and worker adult or pupal offspring, indicating that skew among female eggs reflected skew among daughter queens. On average, both skew among female eggs (measured by the B index) and queen–queen relatedness proved to be low (means±SE=0.06±0.02 and 0.28±0.08, respectively). However, contrary to current skew models, there was no significant association of skew with either relatedness or worker number (used as a measure of productivity). In L. acervorum, predictions of the concession model of skew may hold between but not within populations because queens are unable to assess their relatedness to other queens within colonies. Additional phenomena that may help maintain low skew in the study population include indiscriminate infanticide in the form of egg cannibalism and split sex ratios that penalize reproductive monopoly by single queens within polygynous colonies.     

Selection on social traits in greater spear-nosed bats, Phyllostomus hastatus

Many studies assume that selection molds social traits and have investigated the manner in which this occurs, yet very few studies have measured the strength of selection on social traits in natural populations. In this paper, I report results of phenotypic selection analyses on two social traits – the size of social groups and the frequency of group foraging – in Phyllostomus hastatus, the greater spear-nosed bat. I found significant positive directional selection on individual group foraging frequency, but no directional selection on individuals in different-sized social groups. These results have implications for the structure of social groups, cooperative behavior among group mates, and maternal investment strategies. I argue that combining studies of natural selection on wild populations with experiments to identify the agents and mechanisms of selection can do much to increase our understanding of social evolution.     

Male approach and female avoidance as mechanisms of population discrimination in sagebrush lizards

Reproductive isolation and speciation can result from female choice for particular males. Isolation can also result, however, from male mating preferences or from aggressive encounters which then influence mating decisions. In this study, we use laboratory discrimination trials to study the behavioral mechanisms of population discrimination in sagebrush lizards (Sceloporus graciosus). We specifically ask three questions about population-level discrimination: (1) Does it vary in strength in relation to the geographic distance between the populations? (2) Is it more apparent in inter- or intra-sexual interactions? (3) Does it take the form of attraction or avoidance? We ran 890 trials that tested the ability of male and female sagebrush lizards from one population to discriminate their own population from four other populations. In addition, we utilized both sequential and simultaneous-choice designs, which enabled us to distinguish between attraction and avoidance. We found that most population-level discrimination was exhibited by male lizards preferring to associate with particular types of females, as well as female avoidance of particular types of males. The strength and direction of both discriminations depended on the populations compared and on whether the tests were conducted as sequential- or simultaneous-choice tests, producing a complex relationship between geographic distance and behavioral discrimination. Our results suggest that there are roles for male attraction and female avoidance in population discrimination, reproductive isolation, and speciation.     

Plasma levels of androgens and gonadal development of breeders and helpers in the bell miner (Manorina melanophrys)

Plasma concentration of androgens and gonad development were studied in wild cooperatively breeding bell miners (Manorina melanophrys) of different age, sex and social status. Plasma levels of androgens increased with age of birds. For sexually mature (SM) bell miners, androgen levels were higher in breeders than in nonbreeding (NB) helpers (P < 0.005), and they were marginally higher in females than in males (P < 0.10). Thus female breeders showed the highest plasma levels of androgens. This pattern may be explained by an asymmetry between the sexes in the levels of competition needed to reach and defend breeding status; the degree of intra-sexual competition seems to be higher among female bell miners than among males. In general, male breeders had larger gonads than male SMNB helpers; male breeder gonads produced spermatozoa whereas most helpers' gonads did not. However, old (i.e. 34 months) SMNB male helpers had large gonads producing spermatozoa in spite of having low plasma levels of androgens. We suggest that young male bell miners may be under a voluntary mechanism of reproductive suppression perhaps favoured by inbreeding avoidance; although some aggressive interactions observed between very young helpers (about 2 months) and adult members of the breeding unit suggests that a certain degree of imposition also occurs at young ages. Older male helpers may avoid aggression with the male breeder and imposed reproductive suppression, hence the low circulating levels of androgens, but they have the potential to sneak extra-pair copulations. Gonad size decreased during the non-breeding season in both sexes.