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.     

藻类光竞争模型构建及水体紊动对竞争的影响

光是藻类进行光合作用的重要能量来源,决定藻类的初始生产力,当多种浮游藻类共同生存时,会一起竞争光强.光竞争能力强的浮游藻类浓度增加,光竞争能力较弱的则减少,从而完成水体中浮游藻类优势藻的替代.本文通过在以往建立的单藻模型的基础上,耦合入竞争模型,构建两种（多种）浮游藻类光竞争模型,尝试通过数值模拟,研究不同水体紊动中竞争藻属分别取得竞争优势的机理.模拟结果表明：稳态水体中,竞争藻属的自身垂向迁移决定藻浓度的垂向分布,具有浮力调节能力的微囊藻属占据上层水体优势地位,并取得竞争优势;动态水体中,竞争藻属垂向分布均较为均匀,生长速率决定藻的竞争优势,小球藻属生长速率快,最终易取得竞争优势.此外,考虑浮游藻属的动态自遮蔽,会加速稳态水体中微囊藻属取得竞争优势,并限制动态水体中小球藻属的无限制生长,使得模拟结果更加符合实际情况.     

城市景观水体中浮萍对水华的影响分析

为探究城市景观水体中浮萍对水华暴发的影响,于2015年3—9月在4处典型的景观河道进行采样点布设和检测分析。现场观测从2015年4月2日开始,各采样点陆续出现浮萍,且随着时间的延长,发现浮萍和水华是可以同时存在的;再将检测数据与2014年未出现浮萍生长覆盖的数据进行对比分析可知:浮萍对藻类的生长产生了竞争抑制,推迟了水华的暴发时间,减小了水华的暴发次数;对造成该结果的原因进行显著性分析可得:浮萍对藻类的抑制作用不主要表现在光照的遮挡,而主要是对营养盐的竞争,在有大量的浮萍生长覆盖时,TP成为了限制藻类生长的因素,而TN并不是藻类生长的限制性因素。     

Male moths undertake both pre- and in-copulation mate choice based on female age and weight

Variation in reproductive potential usually occurs among individuals of both sexes; for example, some individuals may carry more gametes or be able to continue to generate more gametes in their reproductive life than others. Therefore, to maximize their reproductive success, both sexes are expected to show adaptations for mate choice. However, most authors concentrated on how females choose their mates with the belief that females invest more in reproduction than males, and males are generally eager to pair with any female. In this paper, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), with special emphasis on male pre- and in-copulation mate choice in relation to the reproductive quality and re-copulation potential of females. We show that in E. kuehniella, younger and heavier females have significantly higher reproductive value than older and lighter ones, and male sperm supply significantly decreases over successive copulations. Males exercise pre-copulation mate choice by selecting females with higher reproductive potential for copulation and in-copulation mate choice by allocating more sperm to females with higher reproductive potential. However, high-quality females are more likely to re-copulate than low-quality ones, and allocation of more sperm to females by males does not increase female reproductive outputs. It is suggested that the allocation of more sperm to high-quality females functions to increase sperm competitiveness against rivals.     

Females go where the food is: does the socio-ecological model explain variation in social organisation of solitary foragers?

The socio-ecological model (SEM) links ecological factors with characteristics of social systems and allows predictions about the relationships between resource distribution, type of competition and social organisation. It has been mainly applied to group-living species but ought to explain variation in social organisation of solitary species as well. The aim of this study was to test basic predictions of the SEM in two solitary primates, which differ in two characteristics of female association patterns: (1) spatial ranging and (2) sleeping associations. Beginning in August 2002, we regularly (re-)captured and marked individuals of sympatric populations of Madame Berthe's and grey mouse lemurs (Microcebus berthae, Microcebus murinus) in Kirindy Forest (Madagascar). We recorded data on spatial patterns, feeding and social behaviour by means of direct observation of radio-collared females. The major food sources of M. berthae occurred in small dispersed patches leading to strong within-group scramble competition and over-dispersed females with a low potential for female associations. In contrast, M. murinus additionally used patchily distributed, high-quality (large) resources facilitating within-group contest competition. The combined influence of less strong within-group scramble and contest as well as between-group contest over non-food resources allowed females of this species to cluster in space. Additionally, we experimentally manipulated the spatial distribution of food sources and found that females adjusted their spatial patterns to food resource distribution. Thus, our results support basic predictions of the SEM and demonstrated that it can also explain variation in social organisation of solitary foragers.