Effects of kinship on territorial conflicts among groups of lions,<Emphasis Type="Italic"> Panthera leo</Emphasis>

Inclusive fitness theory predicts that cost of tolerant behaviour during competitive interactions is lower for relatives than for nonrelatives. Many studies have examined the effect of relatedness on behaviour within social groups. In contrast, kin selection acting among groups has received less attention. The genetic structure of African lion (Panthera leo) populations creates a strong possibility that kin selection among groups modifies behaviour during group conflicts. We used playback experiments and genetic data to investigate the importance of relatedness during simulated territorial disputes in lions. However, we found no effect of relatedness on territorial behaviour. Degree of relatedness did not affect the decision to approach simulated intrusions, nor did it affect the behaviour during approaches. The decision to approach was instead affected by position within the territory and consecutive playback number (a measure of habituation). For playbacks that did elicit an approach, the speed of response was not detectably affected by relatedness, but was affected by odds (the ratio of residents to intruders), number of intruders, number of bouts, presence of cubs, position within the territory, temperature and playback number. Although responses were unaffected by relatedness, it remains possible that other aspects of behaviour during natural encounters among prides are affected by kin selection.Communicated by L. Sterck     

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.     

Asymmetric larval competition in the parasitoid wasp <Emphasis Type="Italic">Nasonia vitripennis</Emphasis>: a role in sex allocation?

Sex allocation theory offers excellent opportunities for testing how animals adjust their behaviour in response to environmental conditions. A major focus has been on instances of local mate competition (LMC), where female-biased broods are produced to maximise mating opportunities for sons. However, the predictions of LMC theory can be altered if there is both local competition for resources during development and an asymmetry between the competitive abilities of the sexes, as has been seen in animals ranging from wasps to birds. In this paper, we test the extent to which asymmetric larval competition alters the predictions of LMC theory in the parasitoid wasp Nasonia vitripennis. We found that the body size of both sexes was negatively correlated with the number of offspring developing within the host. Further, we found that when faced with high levels of competition, the body size of females, but not males, was influenced by the sex ratio of the competing offspring; females were smaller when a higher proportion of the brood was female. This asymmetric competition should favour less biased sex ratios than are predicted by standard LMC theory. We then develop a theoretical model that can be parameterised with our data, allowing us to determine the quantitative consequences of the observed level of asymmetric larval competition for sex allocation. We found that although asymmetric competition selects for less biased sex ratios, this effect is negligible compared to LMC. Furthermore, a similar conclusion is reached when we re-analyse existing data from another parasitoid species where asymmetric larval competition has been observed; Bracon hebetor. Consequently, we suspect that asymmetric larval competition will have its greatest influence on sex ratio evolution in species that have smaller clutches and where local mate competition is not an issue, such as birds and mammals.     

Male signaling behavior and sexual selection in a wolf spider (Araneae: Lycosidae): a test for dual functions

Male signaling behaviors are often studied in a single context but may serve multiple functions (e.g., in male–male competition and female mate choice). We examined the issue of dual function male signals in a wolf spider species Schizocosa ocreata (Hentz) that displays the same species-specific signaling behaviors in both male–male and male–female contexts. These signaling behaviors have been described as either aggression or courtship according to the context observed. We tested the possibility of dual functions by comparing the relationship between behaviors and outcome of male–male contests (winner/loser) and male–female mating encounters (mating success). Frequency, rate, and mean duration of signaling behaviors did not vary with outcome of male–male contests, which appears instead to be based upon relative size and body mass. Winners of contests had significantly greater body mass than losers, and greater mass relative to opponents was significantly associated with probability of winning. Overall, signaling rates were much higher in male–female interactions than in male–male contests and were higher for males that successfully mated than for those that did not mate. Mean duration of some male displays was also greater for males that successfully mated. However, male size was not associated with probability of mating. Taken together, results suggest an intersexual selection context for the current function of male signals in these wolf spiders and that increased display vigor is associated with male mating success.     

Proximal traits and mechanisms for biasing paternity in the red flour beetle Tribolium castaneum (Coleoptera: Tenebrionidae)

A comprehensive understanding of sexual selection requires knowledge of the traits and mechanisms responsible for increasing a male’s paternity share (proportion of progeny sired) relative to that of other males mating with the same female. In this study we manipulated by starvation the expression of traits that might influence male paternity share in Tribolium castaneum. We then conducted experiments to examine how male starvation affects male performance during sequential episodes of sexual selection from mating to progeny production, and investigated female control over specific stages by using live vs dead females. Comparison of starved vs fed males revealed that T. castaneum females have control over spermatophore transfer during mating, as live females rejected inseminations by starved (“low quality”) males. None of the measured male copulatory behaviors (leg-rubbing frequency, asymmetry, and percent of time spent rubbing) affected the probability of successful insemination, but the last two were positively associated with male paternity share. Spermatophore positioning within the female reproductive tract was not affected by male treatment (starved/fed), by female treatment (live/dead), or by male copulatory behaviors. Starvation, however, had a dramatic effect on male reproductive physiology, decreasing both accessory gland size and total number of sperms transferred (but not sperm viability in seminal vesicles). In addition, females who mated to starved males stored fewer sperms in their spermathecae, which, together with decreased ejaculate size, may explain the reduced paternity share of starved males compared to fed males. This study elucidates some cryptic mechanisms influencing male reproductive success and aids our understanding of trait evolution through sexual selection.     

Male influence on sex allocation in the parasitoid wasp Nasonia vitripennis

Sex allocation is an important reproductive decision for parents. However, it is often assumed that females have substantial control over sex allocation decisions, and this is particularly true in haplodiploid insects, in which females apparently determine sex by deciding whether to fertilise an egg (and produce a diploid daughter) or not (and produce a haploid son). Mechanisms by which males may influence sex allocation are not so straightforward, and their potential influence on sex ratios has been somewhat neglected. Here, we test whether males influence offspring sex ratios in the parasitoid wasp Nasonia vitripennis. We show that some of the variation in observed sex ratios can be attributed to males when comparing the affect of male strain on sex ratio. We did not find among-male variation in sex ratio with a less powerful experiment using males from only one strain or an effect of male mating environment. Our data suggest that males can influence female sex ratios and contribute to the variation around the sex ratios optimal for females. However, the influence is not large, suggesting that females have more influence on sex allocation than do males. We conclude by considering whether male influences on sex ratio represent differences in male reproductive competence or deliberate attempts by males to increase their fitness by influencing daughter production.     

The reproductive dilemmas of queenless red dwarf honeybee (Apis florea) workers

Honeybee (Apis) workers cannot mate, but retain functional ovaries. When colonies have lost their queen, many young workers begin to activate their ovaries and lay eggs. Some of these eggs are reared, but most are not and are presumably eaten by other workers (worker policing). Here we explore some of the factors affecting the reproductive success of queenless workers of the red dwarf honeybee Apis florea. Over a 2-year period we collected 40 wild colonies and removed their queens. Only two colonies remained at their translocated site long enough to rear males to pupation while all the others absconded. Absconding usually occurred after worker policing had ceased, as evidenced by the appearance of larvae. Dissections of workers from eight colonies showed that in A. florea, 6% of workers have activated ovaries after 4 days of queenlessness, and that 33% of workers have activated ovaries after 3 weeks. Worker-laid eggs may appear in nests within 4 days and larvae soon after, but this is highly variable. As with Apis mellifera, we found evidence of unequal reproductive success among queenless workers of A. florea. In the two colonies that reared males to pupation and which we studied with microsatellites, some subfamilies had much higher proportions of workers with activated ovaries than others. The significance of absconding and internest reproductive parasitism to the alternative reproductive strategies of queenless A. florea workers is discussed.     

Individual sex ratios and offspring emergence patterns in a parasitoid wasp, Melittobia australica (Eulophidae), with superparasitism and lethal combat among sons

Since the mating of the parasitoid wasp Melittobia australica occurs on their eclosed hosts, the sex ratio is predicted to follow the local mate competition (LMC) theory. However, while LMC models predict that the sex ratio will increase from female-biased toward a 1:1 ratio with an increase in the number of foundresses, the observed female-biased sex ratios (1–5% males) show little increase in response to an increased foundress number. Lethal combat among adult males may serve as an explanation for this observed phenomenon. Using a microsatellite DNA marker, we first examined the individual sex ratio of two foundresses who had sequentially parasitized the same host. Both foundresses produced an extremely female-biased clutch and the sex ratios of the second foundress were only slightly less biased than that of the first. A small number of sons from both foundresses emerged at a constantly low rate over a prolonged period, resulting in a temporal mixture of emerging males derived from both the foundresses. Second, we conducted a one-on-one arena experiment to examine the combat level in relation to the relatedness of the opponents. Almost all the later-emerging males were killed by previously eclosed males irrespective of whether they were sibs or non-sibs. These results suggest that each foundress should not produce males in a single burst, but continue to produce male eggs at a constantly low rate in order to avoid the high mortality of her own sons by lethal male-male combat. This combat may be one of factors in explaining the extremely female-biased sex ratio even with an increasing foundress number.Communicated by R.F.A. Moritz     

Sperm transfer and storage in relation to sperm competition in Callosobruchus maculatus

This paper examines the underlying mechanisms of sperm competition in the beetle Callosobruchus maculatus (F.) (Coleoptera: Bruchidae). Recently developed mathematical models of sperm competition are combined with an empirical investigation of the processes of sperm transfer and storage. During a single insemination virgin males transfer approximately 46000 sperm, 85% more sperm than females can effectively store in their spermathecae. Many of these sperm remain in the bursa copulatrix where they are apparently rapidly degraded and can therefore play no role in fertilization. The spermatheca (primary site of sperm storage) is filled by a single insemination and sperm are lost from this organ at a constant rate. This rate of sperm loss from the spermatheca is insufficient for sperm mixing (without displacement) or sperm stratification to account for the degree of last male sperm precedence measured as P ₂; the proportion of offspring fathered by the second male to mate reported for this species (P ₂ = 0.83, when two inseminations are separated by 24 h). Models of sperm displacement correctly predict high levels of sperm precedence although the precision of these predictions is limited because the proportion of sperm entering the spermatheca cannot be accurately determined. The results suggested that last male sperm precedence in C. maculatus the result of sperm displacement, although the exact mechanism of displacement (sperm-for-sperm or fluid displacement) remains unknown. Possible constraints imposed by female genital anatomy on sperm displacement are discussed.     

Sperm competition in horseshoe crabs (Limulus polyphemus)

Male horseshoe crabs have two mating tactics. Some males come ashore attached to a female (clasping the posterior margin of the females' carapace with their modified pedipalps) and nest with her on the intertidal portion of the beach during the high tide. Other males come ashore unattached and crowd around nesting couples. Fertilization is external and unattached males that are in contact with a pair, i.e. satellite males, release sperm, so the assumption has been that they are fertilizing eggs. We conducted a paternity analysis to determine the proportion of eggs fertilized by attached and satellite males. Pairs with one satellite were observed during nesting on beaches in Florida and Delaware and their eggs were collected and reared to the late trilobite or first instar horseshoe crab stage. DNA was extracted from these offspring and from each adult (female, attached and satellite male) for use in paternity analysis. A Limulus-specific hypervariable microsatellite locus was identified and primers were constructed to amplify this locus via the polymerase chain reaction (PCR). Genotypes of putative parents and offspring were determined by resolving length variants of these PCR products on acrylamide gels. This allowed us to determine parentage of the offspring. We demonstrate that satellite males fertilized 40% of the eggs on average, attached males fertilized 51% and 4% of the eggs that were laid by the female were fathered neither by the attached male nor by the satellite (and 5% could not be determined unambiguously). There is high variability in the success of satellite males, ranging from 0 to 88%. Part of this variability can be explained by the position of the satellite relative to the attached male. We discuss the mechanics of fertilization and the possible advantages for multiple mating in this species.