Low mating frequency of queens in the stingless bee Scaptotrigona postica and worker maternity of males.

Kin selection models of intracolonial conflict over the maternity of males predict that social hymenopteran workers should favour the production of sons and nephews over brothers when the effective mating frequency (me) of the queen is low (me2. Stingless bees have been used to support these models in that me within the group is considered low and workers are thought often to monopolise the parentage of males. We genetically analysed 20 worker and 20 male pupae from each of 10 colonies of the stingless bee Scaptotrigona postica (= Scaptotrigona aff. depilis) using six microsatellite loci and demonstrate queen monandry in eight nests and apparent low me in the other two. However, four colonies contained an additional matriline, possibly due to queen supersedure (serial polygyny), which complicated their genetic structure. Across colonies, workers were responsible for the maternity of 13% of all males. These data are broadly in agreement with predictions from kin selection theory, though the question remains open as to why workers do not secure a greater share of male maternity in this and other stingless bee species in which workers are more closely related to nephews than brothers.

Paternity frequency and maternity of males in some stingless bee species

In monogynous hymenopteran societies, the number of mates of a queen strongly influences the potential for conflict between workers and queens over the maternity of males. Queens always 'prefer' their own sons to sons of workers, regardless of queen mating frequency. When a queen mates once, workers are more closely related to, and therefore are expected to prefer, their own sons and then sons of sisters to sons of the queen. However, if effective paternity frequency exceeds 2, workers on average should prefer queen-produced males to males produced by their sisters. We studied the queen mating frequency of seven stingless bee species: the Mexican species Scaptotrigona mexicana , S. pectoralis and the Australian species Austroplebeia symei , Trigona clypearis , T. hockingsi , T. mellipes and T. sapiens . We then determined whether males arise from eggs laid by workers or queens in A. symei , T. clypearis , T. hockingsi and T. mellipes . We show that all seven species investigated are most likely singly mated and that the queen dominates reproduction. This indicates that the queen's mating frequency alone does not determine whether workers or the queen produces the males.

Single mating in orchid bees (Euglossa, Apinae): implications for mate choice and social evolution

Neotropical orchid bees (Euglossini) are conspicuously different from other corbiculate bees (Apinae) in their lack of advanced sociality and in male use of acquired odors (fragrances) as pheromone-analogues. In both contexts, orchid bee mating systems, in particular the number of males a female mates with, are of great interest but are currently unknown. To assess female mating frequency in the genus Euglossa, we obtained nests from three species in Mexico and Panama and genotyped mothers and their brood at microsatellite DNA loci. In 26 out of 29 nests, genotypes of female brood were fully consistent with being descended from a singly mated mother. In nests with more than one adult female present, those adult females were frequently related, with genotypes being consistent with full sister-sister (r = 0.75) or mother-daughter (r = 0.5) relationships. Thus, our genetic data support the notions of female philopatry and nest-reuse in the genus Euglossa. Theoretically, single mating should promote the evolution of eusociality by maximizing the relatedness among individuals in a nest. However, in Euglossini this genetic incentive has not led to the formation of eusocial colonies as in other corbiculate bees, presumably due to differing ecological or physiological selective regimes. Finally, monandry in orchid bees is in agreement with the theory that females select a single best mate based on the male fragrance phenotype, which may contain information on male age, cognitive ability, and competitive strength.

The effect of social condition and diet on the frequency of male-male agonistic displays in the field cricket, gryllus bimaculatus /

The frequency and type of agonistic displays involved in male-male encounters should be significantly influenced by the presence of females. Discrete agonistic displays vary in energy expenditure and risk, and therefore should be dependent on available resources. The influence of live females and the scent of females, on the frequency of male agonistic displays was observed in a laboratory terrarium using the field cricket Gryllus bimaculatus. The effect of energy constraints on display frequency was also determined. Half the males were fed a diet high in protein and fet; the other males were fed a lower quality diet, for a 7-11 day period. The frequency of five individual displays and mating frequency were recorded using an Event Recorder and notebook. Each group of males was presented with three experimental conditions, over three days, involving the presence or absence of live females and female scent. The presence of females elicited an increase in all displays except antennation; female scent increased the frequency of antennations, mandible flares and grapples, but to a lesser extent than did live females. The frequency of grapples significantly increased for males fed the high quality diet; however diet did not influence the other displays. The combined influence of diet and condition was significant for mandible flare only. Mating frequency was not influenced by diet. However, the frequency ofthe displays were positively correlated with mating frequency for high quality fed males. Escalated displays involving high costs, such as grapple and mandible flare, increased in frequency when the benefits of winning contests were high in G.bimaculatus. Escalation to grapple behaviour was less evident for males fed the lower quality diet as this imposed energy constraints on high cost displays.

The evolution of mating rates in Pieris napi

In the green-veined white butterfly (Pieris napi), females obtain direct fitness benefits from mating multiply and studies have shown that fitness increases seemingly monotonically with number of matings. The reason is that at mating males transfer a large nutritious gift (a so called nuptial gift) to the females that the females use to increase both their fecundity and lifespan. In addition, if exposed to poor food conditions as larvae, females mature at a smaller size compared to males. Accordingly, it was suggested that smaller females could compensate for their size through nuptial feeding by, for instance, mating more frequently. We did not find any support for that hypothesis. On the contrary, larger females remated sooner and had a higher lifetime number of matings. Neither were smaller females able to compensate in any other way, because singly mated females and multiply mated females suffered to the same extent from their smaller size. This thesis also shows that despite the positive relationship between fitness and number of matings, there is a large variation in female mating frequency in wild populations and about every second female mates only once or twice. This variation is not dependent on how often females get courted by males, because female mating frequency was shown not to be affected by male courtship intensity. Hence, the reason for the low mating frequency could either be that males have evolved the ability to manipulate females to mate at a suboptimal rate as a measure of protection against sperm competition, or alternatively, that female mating rate is suppressed by some costs. Using two selection lines, artificially selected for either a high or a low mating rate, we showed that the variation in mating rate was mainly a female trait because which line the females were from affected their mating rate whereas which line the male was from did not. This implies that females mate at a low rate due to hidden costs or due to constraints. The same study also showed that females with a high "intrinsic" mating rate lived shorter, but only when denied remating. This led us to test the hypothesis that the cost females face is to have the ability to mate at a high rate but the cost is only paid when remating opportunities are scarce. However, we found no support for such an idea, because females with a high intrinsic mating rate held in a cold environment where the butterflies were prevented from flying and feeding did not live shorter. Neither was there an effect of a female’s mating rate on her ability to quickly break down and convert male nutrient gifts into egg material. Female mating rate did, on the other hand, affect dispersal tendency, with low mating rate females being more inclined to fly between different habitats. The underlying reason for this is still to be explored.

Complex sociogenetic organization and reproductive skew in a primitively eusocial sweat bee, Lasioglossum malachurum, as revealed by microsatellites

The sweat bees (Family Halictidae) are a socially diverse taxon in which eusociality has arisen independently numerous times. The obligate, primitively eusocial Lasioglossum malachurum, distributed widely throughout Europe, has been considered the zenith of sociality within halictids. A single queen heads a colony of smaller daughter workers which, by mid-summer, produce new sexuals (males and gynes), of which only the mated gynes overwinter to found new colonies the following spring. We excavated successfully 18 nests during the worker- and gyne-producing phases of the colony cycle and analysed each nest's queen and either all workers or all gynes using highly variable microsatellite loci developed specifically for this species. Three important points arise from our analyses. First, queens are facultatively polyandrous (queen effective mating frequency: range 1–3, harmonic mean 1.13). Second, queens may head colonies containing unrelated individuals (n = 6 of 18 nests), most probably a consequence of colony usurpation during the early phase of the colony cycle before worker emergence. Third, nonqueen's workers may, but the queen's own workers do not, lay fertilized eggs in the presence of the queen that successfully develop into gynes, in agreement with so-called 'concession' models of reproductive skew

Complex sociogenetic organization and the origin of unrelated workers in a eusocial sweat bee, Lasioglossum malachurum

Sweat bees (Halictidae) exhibit great interspecific and intraspecific diversity in their social organisation, yet there is remarkably little information on the sociogenetic organisation of any species. Lasioglossum malachurum is a eusocial sweat bee with an annual lifecycle that exhibits considerable variation in its social organisation across its wide geographic range from northern to southern Europe. We collected all adults from 31 L. malachurum nests at Eichkogl, Austria, near the latitudinal centre of its distribution, and genotyped 148 workers using 5 highly variable microsatellite loci developed for this species. Nests were often queenless (48% of nests) during the second phase of worker activity, when colonies were provisioning the sexual brood. Pedigree reconstruction and estimates of nestmate genetic relatedness demonstrated that nests often (32% of nests) contained alien workers, probably as a result of worker drifting from their natal to a foreign nest. Queen effective mating frequency was variable (harmonic mean m(e) = 1.24), but sometimes high (maximum 2.7). These data demonstrate that nests of L. malachurum do not have a classical eusocial sociogenetic organisation (monogyny, monandry) and thereby pose a challenge to exclusively relatedness based arguments for the evolution of eusociality in the taxon.

Sexual behavior and factors affecting female reproduction in house and field crickets

The objective of this investigation was to clarify the adaptive significance of female sexual behaviours in the house cricket, Acheta domesticus, and the Texas field cricket, Gryllus integer. Experiments were focussed primarily on: nutritional factors affecting female reproductive success; the ontogeny of female sexual behaviours; female mating frequency and progeny production; and the pattern of sperm competition. Reproduction of singly mated female A. domesticus assigned to 3 nutritional regimes was compared . Females fed a vitamin and protein-enriched mouse chow, cannibalistic females, and starved females produced on the average, 513 , 200 and 68 offspring, respectively. Cannibals probably could not obtain the same amounts of essential nutrients as females fed mouse chow. Reabsorption of oocytes was likely the major factor contributing to the decreased reproduction of starved females. In addition, female !. domesticus fed mouse chow, but allowed constant access to males produced 11 times as many offspring than did females fed corn meal. Females fed corn meal probably could not absorb or synthesize enough dietary lipids, thus resulting in poor ovariole growth. Female !. domesticus first mate at an average adult age of 7 days, closely corresponding to when they first exhibit positive phonotaxis. Females mate repeatedly and often consume the externally attached spermatophore. In ~. domesticus, females allowed constant access to males produced significantly more offspring than did single maters. Similarly, doubly mated G. integer females produced more offspring than did single maters. This difference resulted largely from the failure of many single maters to reproduce. Remating by female crickets partly functions in offsetting the possibility of a failed initial mating. Nymph production increased significantly with the time the spermatophore was attached in singly mated ~. domesticus. Spermatophore consumption by the female was not affected by male guarding behaviour, and the interval between mating and eating of the spermatophore may often be shorter than the time required for maximum insemination. Some degree of sperm depletion in singly mated !. domesticus and G. integer may have occurred. The patterns of daily offspring production of singly and multiplymated females suggests that a factor provided by a male during mating stimulates female oviposition and/or egg production. Female crickets also might acquire nutrition from spermatophore consumption, a benefit that is augmented by female multiple mating. The electrophoretic examination of various allozymes in ~. integer did not permit determination of a pattern of sperm competition. However, the possibility of last male sperm predominance is related to male guarding behaviour.

The Influence of Male Diet on Life History Traits of Female Mosquitoes

In the field, mosquitoes characteristically feed on sugars soon after emergence and intermittently during their adult lives. Sugar meals are commonly derived from plant nectar and homopteran honeydew, and without them, adults can only survive for a few days on larval reserves. In addition to sugar, females of most species rely on blood for the initiation and maintenance of egg development; thus their reproductive success depends to some extent on the availability of blood hosts. Males, on the other hand, feed exclusively on sugars. Consequently, their sexual maturation and reproductive success is largely dependent upon access to sugar sources. Plant nectar and homopteran honeydew are the two main sugar sources utilized by mosquitoes in the wild. Previous laboratory studies had shown that differences between nectar sources can affect the survivorship and biting frequency of disease vectoring mosquitoes. However, little is known on how sugar composition influence the reproductive processes in male mosquitoes. Male mosquitoes transfer accessory gland proteins and other hormones to their mates along with sperm during mating. In the female, these seminal fluid constituents exert their influence on reproductive genes that control ovulation and vitellogenesis. The present study tests the hypothesis that the mates of males consuming different sugar meals will exhibit varying levels of induction of vitellogenin (a gene which regulates the expression of egg yolk precursor proteins). Real-time quantitative RT-PCR was used to investigate how each sugar meal indirectly influences vitellogenin mRNA abundance in female Anopheles stephensi following mating. Results indicate that mates of nectar-fed males exhibit 2-fold greater change in vitellogenin expression than the mates of honeydew-fed males. However, this response did not occur in non-blood fed controls. These findings suggest that the stimulatory effect of mating on vitellogenesis in blood meal-reliant (i.e. anautogenous) mosquitoes may only be synergistic in nature. The present study also sought to compare the potential fitness costs of mating incurred by females that do not necessarily require a blood meal to initiate a reproductive cycle (i.e., exhibit autogeny). Females of the facultatively autogenous mosquito, Culex molestus were allowed to mate with males sustained on either nectar or honedyew. Mean lifetime fecundity and survivorship of females under the two different mating regimes were then recorded. Additionally, one-dimensional gel electrophoresis was used to verify the transfer of male accessory gland proteins to the sperm storage organs of females during mating.While there was no significant difference in survival between the test treatments, the mates of nectar-fed males produced 11% more eggs on average than mates of honeydew-fed males. However, additional data are needed to justify the extrapolation of these findings to natural settings. These findings prompt further investigation as the differences caused by diet variation in males may be reflected across other life history traits such as mating frequency and insemination capacity.

Characterization of cuticular hydrocarbons of diploid and haploid males, workers and queens of the stingless bee Melipona quadrifasciata

Males, queens and workers of stingless bees show differences in external morphology, behaviour and roles within a colony. In addition, each individual has a cuticular chemical signature responsible for mutual communication that is essential for maintaining the integrity of the colony. In this paper we characterize the cuticular hydrocarbon composition of newly emerged diploid and haploid males, workers and virgin queens of Melipona quadrifasciata by gas chromatography-mass spectrometry (GC/MS) analysis. This is the first time that the cuticular profile of diploid males in a species of stingless bee has been characterized. We found differences in the cuticular hydrocarbon composition among males, workers and virgin queens, recording both qualitative and quantitative differences among individuals of different phenotypes. However, no compound was found exclusively in diploid males. The cuticular chemical profiles of haploid and diploid males were very similar to those of workers. Moreover, the cuticular lipids of males and workers were significantly different from those of queens. Tricosane, pentacosene-2 and 7-methyl-heptacosane were the compounds responsible for this significant separation. This result correlates with the behavioural and morphological differences among these phenotypes.

EVOLUTION OF DOMINANCE UNDER FREQUENCY-DEPENDENT INTRASPECIFIC COMPETITION IN AN ASSORTATIVELY MATING POPULATION

We study the evolution of higher levels of dominance as a response to negative frequency-dependent selection. In contrast to previous studies, we focus on the effect of assortative mating on the evolution of dominance under frequency-dependent intraspecific competition. We analyze a two-locus two-allele model, in which the primary locus has a major effect on a quantitative trait that is under a mixture of frequency-independent stabilizing selection, density-dependent selection, and frequency-dependent selection caused by intraspecific competition for a continuum of resources. The second (modifier) locus determines the degree of dominance at the trait level. Additionally, the population mates assortatively with respect to similarities in the ecological trait. Our analysis shows that the parameter region in which dominance can be established decreases if small levels of assortment are introduced. In addition, the degree of dominance that can be established also decreases. In contrast, if assortment is intermediate, sexual selection for extreme types can be established, which leads to evolution of higher levels of dominance than under random mating. For modifiers with large effects, intermediate levels of assortative mating are most favorable for the evolution of dominance. For large modifiers, the speed of fixation can even be higher for intermediate levels of assortative mating than for random mating.

High frequency of sex and equal frequencies of mating types in natural populations of the ciliate Tetrahymena thermophila.

In ciliate protists, sex involves the temporary joining of two cells of compatible mating type, followed by meiosis and exchange of gametic nuclei between conjugants. Reproduction is by asexual binary fission following conjugation. For the many ciliates with fixed multiple mating types, frequency-dependent sex-ratio theory predicts equal frequencies of mating types, if sex is common in nature. Here, we report that in natural populations of Tetrahymena thermophila sexually immature cells, indicative of recent conjugation, are found from spring through fall. In addition, the seven mating types occur in approximately equal frequencies, and these frequencies appear to be maintained by interaction between complex, multiple mat alleles and environmental conditions during conjugation. Such genotype-environment interaction determining mating type frequency is rare among ciliates.

Polymorphisms in a desaturase 2 ortholog associate with cuticular hydrocarbon and male mating success variation in a natural population of Drosophila serrata

Elucidating the nature of genetic variation underlying both sexually selected traits and the fitness components of sexual selection is essential to understanding the broader consequences of sexual selection as an evolutionary process. To date, there have been relatively few attempts to connect the genetic variance in sexually selected traits with segregating DNA sequence polymorphisms. We set out to address this in a well-characterized sexual selection system - the cuticular hydrocarbons (CHCs) of Drosophila serrata - using an indirect association study design that allowed simultaneous estimation of the genetic variance in CHCs, sexual fitness and single nucleotide polymorphism (SNP) effects in an outbred population. We cloned and sequenced an ortholog of the D. melanogaster desaturase 2 gene, previously shown to affect CHC biosynthesis in D. melanogaster, and associated 36 SNPs with minor allele frequencies > 0.02 with variance in CHCs and sexual fitness. Three SNPs had significant multivariate associations with CHC phenotype (q-value < 0.05). At these loci, minor alleles had multivariate effects on CHCs that were weakly associated with the multivariate direction of sexual selection operating on these traits. Two of these SNPs had pleiotropic associations with male mating success, suggesting these variants may underlie responses to sexual selection due to this locus. There were 15 significant male mating success associations (q-value < 0.1), and interestingly, we detected a nonrandom pattern in the relationship between allele frequency and direction of effect on male mating success. The minor-frequency allele usually reduced male mating success, suggesting a positive association between male mating success and total fitness at this locus.

Changing resonator geometry to boost sound power decouples size and song frequency in a small insect

Despite their small size, some insects, such as crickets, can produce high amplitude mating songs by rubbing their wings together. By exploiting structural resonance for sound radiation, crickets broadcast species-specific songs at a sharply tuned frequency. Such songs enhance the range of signal transmission, contain information about the signaler's quality, and allow mate choice. The production of pure tones requires elaborate structural mechanisms that control and sustain resonance at the species-specific frequency. Tree crickets differ sharply from this scheme. Although they use a resonant system to produce sound, tree crickets can produce high amplitude songs at different frequencies, varying by as much as an octave. Based on an investigation of the driving mechanism and the resonant system, using laser Doppler vibrometry and finite element modeling, we show that it is the distinctive geometry of the crickets' forewings (the resonant system) that is responsible for their capacity to vary frequency. The long, enlarged wings enable the production of high amplitude songs; however, as a mechanical consequence of the high aspect ratio, the resonant structures have multiple resonant modes that are similar in frequency. The drive produced by the singing apparatus cannot, therefore, be locked to a single frequency, and different resonant modes can easily be engaged, allowing individual males to vary the carrier frequency of their songs. Such flexibility in sound production, decoupling body size and song frequency, has important implications for conventional views of mate choice, and offers inspiration for the design of miniature, multifrequency, resonant acoustic radiators.

Females of a tree cricket prefer larger males but not the lower frequency male calls that indicate large body size

In the tree cricket Oecanthus henryi, females are attracted by male calls and can choose between males. To make a case for female choice based on male calls, it is necessary to examine male call variation in the field and identify repeatable call features that are reliable indicators of male size or symmetry. Female preference for these reliable call features and the underlying assumption behind this choice, female preference for larger males, also need to be examined. We found that females did prefer larger males during mating, as revealed by the longer mating durations and longer spermatophore retention times. We then examined the correlation between acoustic and morphological features and the repeatability of male calls in the field across two temporal scales, within and across nights. We found that carrier frequency was a reliable indicator of male size, with larger males calling at lower frequencies at a given temperature. Simultaneous playback of male calls differing in frequency, spanning the entire range of natural variation at a given temperature, revealed a lack of female preference for low carrier frequencies. The contrasting results between the phonotaxis and mating experiments may be because females are incapable of discriminating small differences in frequency or because the change in call carrier frequency with temperature renders this cue unreliable in tree crickets. (C) 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.