Cryptic female choice

The fiddler crab Uca mjoebergi mates both underground in male-defended burrows and on the surface near female-defended burrows. The reproductive tract of Uca species facilitates last-male precedence, suggesting that males that do not guard-mated females are likely to gain very little paternity if the female re-mates with another male. Here, we test the reproductive success of burrow and surface matings using paternity analysis. We found that 100 % of the females that mated in burrows extruded a clutch of eggs. Furthermore, we show conclusively, for the first time in a fiddler crab species, that last-male sperm precedence results in the majority of the female's eggs being fertilised by the burrow-mated male. In contrast, surface matings resulted in significantly fewer females extruding eggs (5.6 %). Paternity analysis also revealed that more than half of the clutches from burrow-mated females showed low levels of extra-pair paternity from previous matings. Although multiple matings appear common in U. mjoebergi, burrow-mated males that guard females are guaranteed a successful mating with extremely high rates of assured paternity. Surface matings therefore appear to be an opportunistic tactic that may increase male reproductive success in a highly competitive environment.

The fiddler crab Uca mjoebergi mates both underground in male-defended burrows and on the surface near female-defended burrows. The reproductive tract of Uca species facilitates last-male precedence, suggesting that males that do not guard-mated females are likely to gain very little paternity if the female re-mates with another male. Here, we test the reproductive success of burrow and surface matings using paternity analysis. We found that 100 % of the females that mated in burrows extruded a clutch of eggs. Furthermore, we show conclusively, for the first time in a fiddler crab species, that last-male sperm precedence results in the majority of the female's eggs being fertilised by the burrow-mated male. In contrast, surface matings resulted in significantly fewer females extruding eggs (5.6 %). Paternity analysis also revealed that more than half of the clutches from burrow-mated females showed low levels of extra-pair paternity from previous matings. Although multiple matings appear common in U. mjoebergi, burrow-mated males that guard females are guaranteed a successful mating with extremely high rates of assured paternity. Surface matings therefore appear to be an opportunistic tactic that may increase male reproductive success in a highly competitive environment.

In polyandrous insects, postcopulatory sexual selection is a pervasive evolutionary force favouring male and female traits that allow control of offspring paternity. Males may influence paternity through adaptations for sperm competition, and females through adaptations facilitating cryptic female choice. Yet, the mechanisms are often complex, involving behaviour, physiology or morphology, and they are difficult to identify. In red flour beetles (Tribolium castaneum), paternity varies widely, and evidence suggests that both male and female traits influence the outcome of sperm competition. To test the role of spermathecal morphology and of sperm storage processes on the outcome of sperm competition, we mated each of 26 virgin females with two males, one of which carrying a phenotypic marker to assign offspring paternity. We manipulated the interval between mating with the first and the second male, to create different conditions of sperm storage (overlapping and nonoverlapping) in the female reproductive tract. To investigate the role of sperm storage more closely, we examined the relationship between paternity and spermathecal morphology in a subset of 14 experimental females. In addition, we also characterized variation in spermathecal morphology in three different strains, wildtype, Chicago black and Reindeer. No significant influence of the intermating interval was found on the paternity of the focal male, although the direction of the difference was in the expected direction of higher last male paternity for longer intervals. Moreover, paternity was not significantly associated with spermathecal morphology, although spermathecal volume, complexity, and tubule width varied significantly and substantially among individuals in all investigated strains.

This article examines the egg hatching periods and environmental and anthropogenic dependencies of five spider species native to North America: the California tarantula (Aphonopelma californicum), the Desert Blonde Tarantula (Aphonopelma chalcodes), the Black Widow (Latrodectus mactans), Cellar Spider (Pholcus phalangioides), and the Brown Recluse (Loxosceles reclusa). A combination of climatic factors, latitude, and anthropogenic influence was found to significantly affect both the timing of egg hatching and the potential for egg dormancy. The study highlights variability in hatching duration and the ecological plasticity of spider reproduction in relation to environmental pressures. Egg dormancy, a delay in hatching due to unsuitable conditions, was documented in multiple species, indicating a strategic evolutionary adaptation.

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Female promiscuity has broad implications for individual behaviour, population genetics and even speciation. In the field cricket Gryllus bimaculatus, females will mate with almost any male presented to them, despite receiving no recorded direct benefits. Previous studies have shown that female crickets can benefit from polyandry through increased hatching success of their eggs. There is evidence that this effect is driven by the potential of polyandrous females to avoid fertilizing eggs with sperm from genetically incompatible males. We provide direct evidence supporting the hypothesis that polyandry is a mechanism to avoid genetic incompatibilities resulting from inbreeding. Using microsatellite markers we examined patterns of paternity in an experiment where each female mated with both a related and an unrelated male in either order. Overall, unrelated males were more successful in gaining paternity than were related males, but this effect was driven by a much greater success of unrelated males when they were the first to mate.

Male and female social roles are largely predicated on the fact that male and female reproductive functions are separated in different individuals. This paper asks why gonochorism rather than hermaphroditism, is the rule among vertebrates. We argue that hermaphroditism may be unstable in the face of heterogeneity. Building on the Bateman principle -access to eggs, not sperm, limits reproductive success -and in line with Trivers-Willard, we show that low quality individuals will prefer to be all female. Moreover, without secondary sexual differentiation (SSD), males cannot exist in equilibrium. With sufficient SSD, however, males may outcompete hermaphrodites. As a result, while hermaphrodites may coexist with males and females, they mate among themselves only. The lack of interbreeding between hermaphrodites and gonochorists may form the basis for further speciation. Furthermore, while hermaphrodites strive to mate their male function and preserve their female function, equilibrium hermaphroditic mating is reciprocal. Reciprocal mating, in turn, makes hermaphrodites vulnerable to male-to-male violence, a form of SSD that may have contributed to the rarity of hermaphroditism.

Both males and females of the spider Leiicaiige mariana (Taczanowski 1881) contribute material to the plugs that often occlude the genital openings of females in the field. Males were sometimes unable to remove or penetrate these plugs, but overcame others using three different mechanical mechanisms: snag the plug and pull it off; break and penetrate through it; and break its adhesion to the epigynum by injecting material under it. They used their genitalia to accomplish these tasks, despite the fact that the genital bulb lacks muscles and innervation, thus limiting the male's ability to guide genital movements precisely. The effects of two male genital structures, the conductor tip and the conductor hook on sperm transfer and genital plug removal were tested by direct observations of their morphology and behavior, and by experimental removal of structures from one but not the other palp of the same male. Removal of the conductor tip reduced sperm transfer, while removal of both the hook and the conductor reduced plug removal. A preliminary characterization of palp movements and their sequences did not reveal any behavior that seemed especially designed for removing plugs, as opposed to inseminating the female.

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Males. of two species of Xyonysius court females both before and during copulation. Courtship during copulation ("copulatory courtship") is more elaborate, and differs between the two species. Male genital sclerites are moved in ways suggesting that they stimulate females during copulation. It is argued that copulatory courtship may be under sexual sclcction by cryptic female choice.

The male genitalia of many animal groups have elaborate and species-specific forms. One hypothesis to explain why this is so is that male genitalia function as stimulatory devices that are under sexual selection by cryptic female choice. This report is based on a videotaped observation of a single male of an unidentified species of Tipula (Bellarina) from San José Province, Costa Rica. A male crane fly stridulated (produced vibrations) during copulation using sustained and stereotyped movements of file and scraper structures on his genitalia. Males of related species have similar file and scraper structures on their genitalia, suggesting that they probably also stridulate during copulation.

Female promiscuity has broad implications for individual behaviour, population genetics and even speciation. In the field cricket Gryllus bimaculatus, females will mate with almost any male presented to them, despite receiving no recorded direct benefits. Previous studies have shown that female crickets can benefit from polyandry through increased hatching success of their eggs. There is evidence that this effect is driven by the potential of polyandrous females to avoid fertilizing eggs with sperm from genetically incompatible males. We provide direct evidence supporting the hypothesis that polyandry is a mechanism to avoid genetic incompatibilities resulting from inbreeding. Using microsatellite markers we examined patterns of paternity in an experiment where each female mated with both a related and an unrelated male in either order. Overall, unrelated males were more successful in gaining paternity than were related males, but this effect was driven by a much greater success of unrelated males when they were the first to mate.

Direct costs and benefits to females of multiple mating have been shown to have large effects on female fecundity and longevity in several species. However, with the exception of studies examining genetic benefits of polyandry, little attention has been paid to the possible effects on offspring of multiple mating by females. We propose that nongenetic effects of maternal matings on offspring fitness are best viewed in the same context as other maternal phenotype effects on offspring that are well known even in species lacking parental care. Hence, matings can exert effects on offspring in the same way as other maternal environment variables, and are likely to interact with such effects. We have conducted a study using yellow dung flies (Scathophaga stercoraria), in which we independently manipulated female mating rate, number of mates and maternal thermal environment and measured subsequent fecundity, hatching success, and offspring life-history traits. To distinguish between direct effects of matings and potential genetic benefits of polyandry we split broods and reared offspring at three different temperature regimes. This allowed us to demonstrate that although we could not detect any simple benefits or costs to matings, there are effects of maternal environment on offspring and these effects interact with female mating regime affecting offspring fitness. Such interactions between female phenotype and the costs and benefits of matings have potentially broad implications for understanding female behavior.

Selection clearly focuses on differences in reproduction, but studies of reproductive physiology generally have been carried out in a near vacuum of modern evolutionary theory. This lack of contact between the two fields may be about to change. New ideas indicate that sexual selection by cryptic female choice has affected the evolution of products in male semen that influence female reproductive behavior and physiology.

Queens of Acromyrmex leaf-cutting ants store sperm of multiple males after a single mating flight, and never remate even though they may live for decades and lay tens of thousands of eggs. Sperm of different males are initially transferred to the bursa copulatrix and compete for access to the long-term storage organ of queens, but the factors determining storage success or failure have never been studied. We used in vitro experiments to show that reproductive tract secretions of Acromyrmex echinatior queens increase sperm swimming performance by at least 50% without discriminating between sperm of brothers and unrelated males. Indiscriminate female-induced sperm chemokinesis makes the likelihood of storage directly dependent on initial sperm viability and thus provides a simple mechanism to secure maximal possible reproductive success of queens, provided that initial sperm motility is an accurate predictor of viability during later egg fertilization.

Copulation behavior has often been shaped by sexually selected sperm competition or cryptic female choice. However, manipulation of previously deposited ejaculates is unknown in the social Hymenoptera and the degree to which sperm competes after insemination or is actively selected by females has remained ambiguous. We studied the mating process in the leaf-cutting ants Atta colombica and A. cephalotes, which belong to one of the few derived social insect lineages where obligate multiple mating has evolved. As copulations often occur at night and in remote places, direct observations were impossible, so we had to reconstruct the sequential copulation events by morphological analysis of the male and female genitalia and by tracking the process of sperm transfer and sperm storage. We show that Atta male genitalia have two external rows of spiny teeth, which fit into a specialized pouch organ in the female sexual tract. Reconstruction of the sperm storage process indicated that sperm is transferred to the spermatheca during or immediately after ejaculation and without being mixed with sperm and seminal fluids from other males. A convergent mechanism of direct sperm transfer to the spermatheca of queens is known from two species of dwarf honeybees. Direct sperm transfer may restrict female control over the sperm storage process and the number of males that contribute to the stored sperm.

Direct costs and benefits to females of multiple mating have been shown to have large effects on female fecundity and longevity in several species. However, with the exception of studies examining genetic benefits of polyandry, little attention has been paid to the possible effects on offspring of multiple mating by females. We propose that nongenetic effects of maternal matings on offspring fitness are best viewed in the same context as other maternal phenotype effects on offspring that are well known even in species lacking parental care. Hence, matings can exert effects on offspring in the same way as other maternal environment variables, and are likely to interact with such effects. We have conducted a study using yellow dung flies (Scathophaga stercoraria), in which we independently manipulated female mating rate, number of mates and maternal thermal environment and measured subsequent fecundity, hatching success, and offspring life-history traits. To distinguish between direct effects of matings and potential genetic benefits of polyandry we split broods and reared offspring at three different temperature regimes. This allowed us to demonstrate that although we could not detect any simple benefits or costs to matings, there are effects of maternal environment on offspring and these effects interact with female mating regime affecting offspring fitness. Such interactions between female phenotype and the costs and benefits of matings have potentially broad implications for understanding female behavior.

the subject of the study is the behaviour of Mallard’s Anas platyrhynchos and was carried out in the city of szczecin (Poland) in 2009–2010. we compared frequency of fights, rapes, courtships, copulations and whistles on a water basins within both big and small flocks, and in different phenology periods. the study shows significant relationship between season, number of ducks on a pond and frequency of fights, as well as between whistles and fights, whistles and season and courtships and season. the results show that the most active period for ducks are the autumn and winter months (pairing period), while more aggressive behaviours can be observed when the population density is high.

Queens of Acromyrmex leaf-cutting ants store sperm of multiple males after a single mating flight, and never remate even though they may live for decades and lay tens of thousands of eggs. Sperm of different males are initially transferred to the bursa copulatrix and compete for access to the long-term storage organ of queens, but the factors determining storage success or failure have never been studied. We used in vitro experiments to show that reproductive tract secretions of Acromyrmex echinatior queens increase sperm swimming performance by at least 50% without discriminating between sperm of brothers and unrelated males. Indiscriminate female-induced sperm chemokinesis makes the likelihood of storage directly dependent on initial sperm viability and thus provides a simple mechanism to secure maximal possible reproductive success of queens, provided that initial sperm motility is an accurate predictor of viability during later egg fertilization.

The sagebrush cricket, Cyphoderris strepitans, is one of only five extant species belonging to an ancient insect lineage, the Haglidae, believed to be ancestral to modern-day crickets and katydids (Orthoptera: Ensifera) (Morris and Gwynne 1978). C. strepitans occurs exclusively in mountainous areas of Wyoming and Colorado, where it is found primarily in high-altitude sagebrush meadow habitat. Adults become sexually active in May, shortly after snowmelt, and remain active for the next 4-6 weeks. Each night of the breeding season, males emerge from the soil litter shortly after sunset, climb into the sagebrush and begin to sing, presumably to attract sexually receptive females. Copulation is initiated when a receptive female climbs onto the dorsum of a male, at which time he attempts to transfer a spermatophore. During copulation, the female feeds on the male's fleshy metathoracic wings and ingests any haemolymph leaking from the wounds she inflicts (Sakaluk et al. 1987, Morris et...

The design and implementation of assisted reproductive technology to improve genetic diversity and augment captive populations is an important but rarely applied research field in reptiles. Using the corn snake (Elaphe gutatta) as a model, the Henry Doorly Zoo recently produced offspring born as a result of artificial insemination using both fresh, diluted semen, and diluted semen stored at refrigeration for 3 days. Semen was collected noninvasively from sexually mature male corn snakes using a gentle massaging technique, extended in medium then inseminated into the oviducts of adult females. Using molecular genetic techniques to confirm or refute the success of the insemination using primers developed for the black rat snake, Elaphe obsolete, all possible parents and offspring genotypes were evaluated. A paternity-by-exclusion analysis verified that the offspring were in fact a result of artificial insemination. Zoo Biol 26:363-369, 2007.

Fertility depends, in part, on interactions between male and female reproductive proteins inside the female reproductive tract (FRT) that mediate postmating changes in female behaviour, morphology, and physiology. Coevolution between interacting proteins within species may drive reproductive incompatibilities between species, yet the mechanisms underlying postmating-prezygotic (PMPZ) isolating barriers remain poorly resolved. Here, we used quantitative proteomics in sibling Drosophila species to investigate the molecular composition of the FRT environment and its role in mediating species-specific postmating responses. We found that (i) FRT proteomes in D. simulans and D. mauritiana virgin females express unique combinations of secreted proteins and are enriched for distinct functional categories, (ii) mating induces substantial changes to the FRT proteome in D. mauritiana but not in D. simulans, and (iii) the D. simulans FRT proteome exhibits limited postmating changes irrespective of whether females mate with conspecific or heterospecific males, suggesting an active female role in mediating reproductive interactions. Comparisons with similar data in the closely related outgroup species D. melanogaster suggest that divergence is concentrated on the D. simulans lineage. Our study suggests that divergence in the FRT extracellular environment and postmating response contribute to previously described patterns of PMPZ isolation and the maintenance of species boundaries.

Genital morphology shows peculiar patterns of variation among insect species. Traditionally, genital species specificity has been assumed to serve as a mechanical isolation system between species (the lock-and-key hypothesis). Most recent studies suggest, however, that such variation may also be because of sexual selection. These two hypotheses give different predictions on genital variation within and between species. We tested the lock-and-key hypothesis by morphometrically exploring variation and allometry in male genitalia in three closely related Euxoa moth species. Single genital characteristics usually show overlap between species. As a whole, internal genitalia distinguish species better than external genitalia. The size of genitalia is generally correlated with body size, but the relationship is strongly negatively allometric so that the size of internal genitalia increases least with body size. These findings support the lock-and-key hypothesis. Both external and internal male genitalia show morphometric variation both within and between species, but the variation is significantly smaller in external genitalia. As internal genitalia are assumed to work as 'keys' in moths, this finding does not support the predictions of the lock-and-key hypothesis. Therefore, we cannot unambiguously support or reject this hypothesis. Our results agree well with the sexual selection hypotheses, particularly, the one-size-fits-all concept of the cryptic female choice hypothesis, which suggests stabilizing selection on genital size, but at the same time allows genital shape to vary relatively considerably.

Insect genitals vary greatly among species and provide a key tool for species-level taxonomy. Insects differing in the genitalia are often treated as discrete, reproductively isolated species. This principle dates back to the lock-and-key hypothesis, which states that genitalia vary between species in order to provide a mechanical reproductive isolation system. Thus, the hypothesis assumes low within-species variability in genital traits. However, recent studies suggest that sexual selection may be responsible for the evolution of insect genitalia. We studied allometry and genital size and shape variation in a dimorphic moth Selenia tetralunaria. We found that the genitalia showed negative allometry in relation to body size as reported in many insect and spider species. This allometry was stronger in internal genital structures than it was in external genitalia. We also found that there was more variation in internal compared with external genitalia. Finally, we found that the shape of genital structures differed between morphs in all three examined areas. S. tetralunaria is among the first reported cases of genitally dimorphic insect species. Considerable variation in internal genitalia and especially the presence of genital shape differences between morphs were not consistent with the predictions of the lock-and-key hypothesis.

Postcopulatory sexual selection is credited as a principal force behind the rapid evolution of reproductive characters, often generating a pattern of correlated evolution between interacting, sex-specific traits. Because the female reproductive tract is the selective environment for sperm, one taxonomically widespread example of this pattern is the co-diversification of sperm length and female sperm-storage organ dimension. In Drosophila, having testes that are longer than the sperm they manufacture was believed to be a universal physiological constraint. Further, the energetic and time costs of developing long testes have been credited with underlying the steep evolutionary allometry of sperm length and constraining sperm length evolution in Drosophila. Here, we report on the discovery of a novel spermatogenic mechanism—sperm cyst looping—that enables males to produce relatively long sperm in short testis. This phenomenon (restricted to members of the saltans and willistoni species...

We evaluated the in£uence of pre-and post-copulatory sexual selection upon male reproductive traits in a naturally promiscuous species, Drosophila melanogaster. Sexual selection was removed in two replicate populations through enforced monogamous mating with random mate assignment or retained in polyandrous controls. Monogamous mating eliminates all opportunities for mate competition, mate discrimination, sperm competition, cryptic female choice and, hence, sexual con£ict. Levels of divergence between lines in sperm production and male ¢tness traits were quanti¢ed after 38^81 generations of selection. Three a priori predictions were tested: (i) male investment in spermatogenesis will be lower in monogamy-line males due to the absence of sperm competition selection, (ii) due to the evolution of increased male benevolence, the ¢tness of females paired with monogamy-line males will be higher than that of females paired with control-line males, and (iii) monogamy-line males will exhibit decreased competitive reproductive success relative to control-line males. The ¢rst two predictions were supported, whereas the third prediction was not. Monogamy males evolved a smaller body size and the size of their testes and the number of sperm within the testes were disproportionately further reduced. In contrast, the ¢tness of monogamous males (and their mates) was greater when reproducing in a non-competitive context: females mated once with monogamous males produced o¡spring at a faster rate and produced a greater total number of surviving progeny than did females mated to control males. The results indicate that sexual selection favours the production of increased numbers of sperm in D. melanogaster and that sexual selection favours some male traits conferring a direct cost to the fecundity of females.

Postmating sexual selection theory predicts that in allopatry reproductive traits diverge rapidly and that the resulting differentiation in these traits may lead to restrictions to gene flow between populations and, eventually, reproductive isolation. In this paper we explore the potential for this premise in a group of damselflies of the family Calopterygidae, in which postmating sexual mechanisms are especially well understood. Particularly, we tested if in allopatric populations the sperm competition mechanisms and genitalic traits involved in these mechanisms have indeed diverged as sexual selection theory predicts. We did so in two different steps. First, we compared the sperm competition mechanisms of two allopatric populations of Calopteryx haemorrhoidalis (one Italian population studied here and one Spanish population previously studied). Our results indicate that in both populations males are able to displace spermathecal sperm, but the mechanism used for sperm removal between both populations is strikingly different. In the Spanish population males seem to empty the spermathecae by stimulating females, whereas in the Italian population males physically remove sperm from the spermathecae. Both populations also exhibit differences in genital morphometry that explain the use of different mechanisms: the male lateral processes are narrower than the spermathecal ducts in the Italian population, which is the reverse in the Spanish population. The estimated degree of phenotypic differentiation between these populations based on the genitalic traits involved in sperm removal was much greater than the differentiation based on a set of other seven morphological variables, suggesting that strong directional postmating sexual selection is indeed the main evolutionary force behind the reproductive differentiation between the studied populations. In a second step, we examined if a similar pattern in genital morphometry emerge in allopatric populations of this and other three species of the same family (Calopteryx splendens, C. virgo and Hetaerina cruentata). Our results suggest that there is geographic variation in the sperm competition mechanisms in all four studied species. Furthermore, genitalic morphology was significantly divergent between populations within species even when different populations were using the same copulatory mechanism. These results can be explained by probable local coadaptation processes that have given rise to an ability or inability to reach and displace spermathecal sperm in different populations. This set of results provides the first direct evidence of intraspecific evolution of genitalic traits shaped by postmating sexual selection.

Odonates (dragonflies) are well known for the ability of the males to displace sperm stored in the female's spermstorage organs during copulation. By this means, copulating males are able to increase their fertilization success. This ability has been used as an example to illustrate a conflict of interests between the sexes in which males have evolved sperm-displacement mechanisms whilst females have presumably evolved means to avoid sperm displacement. The present review has four aims: (1) to describe the copulatory mechanisms used during sperm displacement; (2) to analyse the causes of sperm usage patterns; (3) to discuss this information using current hypotheses on conflict between the sexes; (4) to illuminate topics for further research. Four copulatory mechanisms are described: sperm removal (physical withdrawal of stored sperm), sperm repositioning ('pushing' of rival sperm to sites where its use will be least likely), female sensory stimulation to induce sperm ejection, and sperm flushing (displacement of sperm using the copulating male's sperm). Sperm-precedence studies in Odonata are scarce and their values vary considerably between species. In those species in which sperm displacement is incomplete, the last copulating male obtains a high but variable short-term fertilization success which decreases with time. Some male and female factors affecting sperm precedence patterns are mentioned: (1) male variation in genital morphology; (2) duration of copulation influenced by the male (the longer the copulation, the more stored sperm displaced); (3) adaptations of the sperm-storage organs that allow the female to manipulate the sperm she has received (i.e. avoiding sperm displacement, redistributing sperm masses, favouring sperm located in certain sites and ejecting sperm after copulation). We suggest that male and female odonates have co-evolved at the level of genital function with the control of stored sperm as the focus of the conflict. The benefits for males in this co-evolution lie in maximizing their fertilization success. However, it is not clear what females obtain from storing sperm and making it unreachable during sperm displacement. Two hypothetical benefits that females may obtain for which some evidence has been gathered are genetic diversity and viability genes. It is finally suggested that odonates can become excellent subjects of study for testing current ideas related to sexual conflict and speciation processes through sexual selection.

Bilateral asymmetry in the genitalia is a rare but widely dispersed phenomenon in the animal tree of life. In arthropods, occurrences vary greatly from one group to another and there seems to be no common explanation for all the independent origins. In spiders, genital asymmetry appears to be especially rare. Few examples have been studied in detail but isolated reports are scattered in the taxonomic literature. Based on a broad literature study, we found several species in thirteen families with evidence of genital asymmetry, mostly expressed only in females. Our review suggests that spider genital asymmetries, although rare, are more common than previously thought and taxonomic descriptions and illustrations are a useful but not entirely reliable tool for studying them. Here we also document thoroughly the case of the liocranid spider Teutamus politus. We collected live specimens to observe male-female interactions and document their genital morphology. We consider T. politus to be the first known case of directional asymmetry and the first report of developmentally asymmetric male genitals in Entelegynae spiders. Generalities, evolution and categorization of asymmetry in spiders are further discussed.

This is particularly apparent within the insects that show high variations in genitalic form between closely related species. Static allometry is one of the effective approaches for quantification of such variation. Despite the crucial roles of the parasitic Hymenoptera in the ecosystems, little is known about the sexual selection in this order, compared with other insect orders. We examined the allometry of different morphological traits in 35 males and 35 females of Habrobracon hebetor (Say, 1836) (Hymenoptera: Braconidae) from a laboratory colony reared in Mashhad city, for the first time. The aim was the investigation of allometric relationships of different body traits as a way for quantification of the natural selection impacts on the different body parts. 12 genitalic and non-genitalic body parts of H. hebetor males and females were photographed and measured. Principal Component Analysis (PCA) was used to explore the variance of the traits and two regression analysis methods to obtain the allometric slopes. All the non-genitalic traits in male and female wasps showed isometry except pterostigma width in male wasps which showed positive allometry. In male genitalia, two traits showed strong negative allometry and one trait showed isometry. Our findings showed that in this species males with an average size of genitalia were more successful in generating viable offspring than males with relatively smaller or larger genitalia sizes and this is irrespective of the overall body size. Our results showed for the first time that such stabilizing sexual selection might operate on genital size in the braconid wasps.

The pre-maturation social environment experienced by females may affect their post-maturation reproductive strategies, including mating preferences and investment in offspring. Whether the pre-maturation social environment also affects other aspects of females’ reproductive strategies, such as the degree of polyandry and post-copulatory decisions, is still an open question. To address this question, we performed laboratory experiments using the cricket Teleogryllus commodus, a highly polyandrous species. Previous studies showed that juvenile females reared in an acoustic environment with recorded male calls with different rates (variable-quality, VQ) are more responsive to high-quality calls than females reared in an environment with recorded male calls with only high rates (high-quality, HQ). We exposed juvenile females to these same two acoustic environments and estimated their degree of polyandry, offspring fitness, and time of spermatophore retention. We found that the juvenile ...

Gonipterus scutellatus es un escarabajo fitófago que se alimenta principalmente de las diferentes especies de Eucalyptus spp. Hasta 2012 se consideraba una sola especie, pero nuevos datos indicaron que en realidad es un complejo, denominado "scutellatus", de 10 especies crípticas con diferentes niveles de preferencia sobre eucalipto. Este complejo es originario de Australia, y contiene varias especies invasoras, tales como: G. platensis, G. pulverulentus y Gonipterus sp. n. 2, siendo estas una de las principales plagas del eucalipto fuera de su región de origen. Las hembras ponen alrededor de 270 huevos en su vida, agrupados en coprotecas de 8 a 10 huevos. Este insecto experimenta una metamorfosis completa, y el estado larval tiene 4 instares, y su ciclo de vida reporta gran variación con una duración, entre 42.7 a 255.95 días debido a la temperatura y la dieta. Los estudios sobre la preferencia alimenticia y la susceptibilidad de las especies de Eucalyptus muestran que G. platensis y G. pulverulentus tienen preferencias similares, y las especies del subgénero Symphiomyrtus son las más afectadas por esta plaga. Palabras clave: Gonipterus, ciclo de vida, Eucalyptus, revisión Revisión de la morfología, biología e historia de vida del complejo Gonipterus scutellatus (COLEOPTERA: Curculionidae) Review of the morphology, biology and life history of the Gonipterus scutellatus complex (COLEOPTERA: Curculionidae

Body size is commonly associated with biological features such as reproductive capacity, competition, and resource acquisition. Many studies have tried to understand how these isolated factors can affect the body pattern of individuals. However, little is known about how interactions among species in multitrophic communities determine the body shape of individuals exploiting the same resource. Here, we evaluate the effect of fruit infestation, parasitism rate, and seed biomass on size, allometric and asymmetric patterns of morphological structures of insects that exploit the same resource. To test it, we measured 750 individuals associated with the plant Senegalia tenuifolia (Fabaceae), previously collected over three consecutive years. Negative allometry was maintained for all species, suggesting that with increasing body size the body structure did not grow proportionally. Despite this, some variations in allometric slopes suggest that interactions in a multitrophic food web can s...

Why dominant males experiencing intense sperm competition sometimes show low investments in sperm production is not always obvious. One well-documented example is that of the external fertilizing teleost, the Arctic charr (Salvelinus alpinus), where individuals becoming dominant reduce sperm production and sperm swimming speed in water compared to subordinates. Here, we report how ovarian fluid differentially influences sperm velocity of dominant and subordinate male Arctic charr. That is, sperm from dominant males increase their velocity in water diluted ovarian fluid compared to that observed in water, while sperm from subordinates, on the other hand, decrease velocity in ovarian fluid compared to that observed in water. Thus, subordinates, who invest more resources in their sperm and usually show the highest sperm velocity in water, have lower gains from their investment than dominant males when sperm are swimming in ovarian fluid. In sum, our result suggests that ovarian fluid increase sperm velocity more in dominant males than in subordinate males. Although this finding could partly be caused by cryptic female choice exerted by the ovarian fluid for sperm from dominant males, an alternative and more parsimonious explanation is that sperm from dominant males may simply be better designed for swimming in ovarian fluid compared to sperm from subordinate males. Thus, sperm production in the two reproductive roles seems to be adaptively tailored to different external environments.

An operant conditioning situation was used to relate the lever-pressing performance of female rhesus monkeys to different measures of social, sexual, and agonistic behavior that underlie the formation and dissolution of consort bonds. Nine females were trained to press a lever 250 times to gain access to a male partner. After access, a standard 60-min behavioral test took place (1,440 tests). Data were analyzed independently of the stage of the menstrual cycle. Eight females were tested with two males, and every female gained access faster with one male (preferred partner). For all eight females, the preferred male was the one that spent more time grooming the female. For five females, the preferred partner was also the one that ejaculated more frequently. For four females, where agonistic interactions with males could be evaluated, the preferred male was the one that elicited fewer submissive behavioral patterns. These results indicate that the operant behavior of female rhesus monkeys was positively reinforced by social and sexual factors and negatively reinforced by agonistic interactions and may thus provide a measure of the strength of consort bonds.

How males and females contribute to joint reproductive success has been a long-standing question in sexual selection. Under postcopulatory sexual selection (PSS), paternity success is predicted to derive from complex interactions among females engaging in cryptic female choice and males engaging in sperm competition. Such interactions have been identified as potential sources of genetic variation in sexually selected traits but are also expected to inhibit trait diversification. To date, studies of interactions between females and competing males have focused almost exclusively on genotypes and not phenotypic variation in sexually selected traits. Here, we characterize within- and between-sex interactions inDrosophila melanogasterusing isogenic lines with heritable variation in both male and female traits known to influence competitive fertilization. We found surprisingly few genotypic interaction effects on various stages of PSS such as female remating interval, copulation duration...

Directional dominance is a prerequisite of inbreeding depression. Directionality arises when selection drives alleles that increase fitness to fixation and eliminates dominant deleterious alleles, while deleterious recessives are hidden from it and maintained at low frequencies. Traits under directional selection (i.e., fitness traits) are expected to show directional dominance and therefore an increased susceptibility to inbreeding depression. In contrast, traits under stabilizing selection or weakly linked to fitness are predicted to exhibit little-to-no inbreeding depression. Here, we quantify the extent of inbreeding depression in a range of male reproductive characters and then infer the mode of past selection on them. The use of transgenic populations of Drosophila melanogaster with red or green fluorescenttagged sperm heads permitted in vivo discrimination of sperm from competing males and quantification of characteristics of ejaculate composition, performance, and fate. We found that male attractiveness (mating latency) and competitive fertilization success (P2) both show some inbreeding depression, suggesting they may have been under directional selection, whereas sperm length showed no inbreeding depression suggesting a history of stabilizing selection. However, despite having measured several sperm quality and quantity traits, our data did not allow us to discern the mechanism underlying the lowered competitive fertilization success of inbred (f = 0.50) males.

How sperm from competing males are used to fertilize eggs is poorly understood yet has important implications for postcopulatory sexual selection. Sperm may be used in direct proportion to their numerical representation within the fertilization set or with a bias toward one male over another. Previous theoretical treatments have assumed a single sperm-storage organ, but many taxa possess multiple organs or store sperm within multiple regions of the reproductive tract. In Drosophila, females store sperm in two distinct storage organ types: the seminal receptacle (SR) and the paired spermathecae. Here, we expand previous "raffle" models to describe "fertilization bias" independently for sperm within the SR and the spermathecae and estimate the fertilization set based on the relative contribution of sperm from the different sperm-storage organ types. We apply this model to three closely related species to reveal rapid divergence in the fertilization set and the potential for female sperm choice.

How females store and use sperm after remating can generate postcopulatory sexual selection on male ejaculate traits. Variation in ejaculate performance traits generally is thought to be intrinsic to males but is likely to interact with the environment in which sperm compete (e.g., the female reproductive tract). Our understanding of female contributions to competitive fertilization success is limited, however, in part because of the challenges involved in observing events within the reproductive tract of internally fertilizing species while discriminating among sperm from competing males. Here, we used females from crosses among isogenic lines of Drosophila melanogaster , each mated to two genetically standardized males (the first with green- and the second with red-tagged sperm heads) to demonstrate heritable variation in female remating interval, progeny production rate, sperm-storage organ morphology, and a number of sperm performance, storage, and handling traits. We then used ...

Postcopulatory sexual selection is credited with driving rapid evolutionary diversification of reproductive traits and the formation of reproductive isolating barriers between species. This judgment, however, has largely been inferred rather than demonstrated due to general lack of knowledge about processes and traits underlying variation in competitive fertilization success. Here, we resolved processes determining sperm fate in twice-mated females, using transgenic Drosophila simulans and Drosophila mauritiana populations with fluorescently labeled sperm heads. Comparisons among these two species and Drosophila melanogaster revealed a shared motif in the mechanisms of sperm precedence, with postcopulatory sexual selection potentially occurring during any of the three discrete stages: (1) insemination; (2) sperm storage; and (3) sperm use for fertilization, and involving four distinct phenomena: (1) sperm transfer; (2) sperm displacement; (3) sperm ejection; and (4) sperm selection for fertilizations. Yet, underlying the qualitative similarities were significant quantitative differences in nearly every relevant character and process. We evaluate these species differences in light of concurrent investigations of within-population variation in competitive fertilization success and postmating/prezygotic reproductive isolation in hybrid matings between species to forge an understanding of the relationship between microevolutionary processes and macroevolutionary patterns as pertains to postcopulatory sexual selection in this group.

Selection to avoid inbreeding is predicted to vary across species due to differences in population structure and reproductive biology. Over the past decade, there have been numerous investigations of postcopulatory inbreeding avoidance, a phenomenon that first requires discrimination of mate (or sperm) relatedness and then requires mechanisms of male ejaculate tailoring and/or cryptic female choice to avoid kin. The number of studies that have found a negative association between male-female genetic relatedness and competitive fertilization success is roughly equal to the number of studies that have not found such a relationship. In the former case, the underlying mechanisms are largely unknown. The present study was undertaken to verify and expand upon a previous report of postcopulatory inbreeding avoidance in D. melanogaster, as well as to resolve underlying mechanisms of inbreeding avoidance using transgenic flies that express a sperm head-specific fluorescent tag. However, siblings did not have a lower fertilization success as compared to unrelated males in either the first (P 1) or second (P 2) mate role in sperm competition with a standard unrelated competitor male in our study population of D. melanogaster. Analyses of mating latency, copulation duration, egg production rate, and remating interval further revealed no evidence for inbreeding avoidance.

Contrary to early predictions of sperm competition theory, postcopulatory sexual selection favoring increased investment per sperm (e.g., sperm size, sperm quality) has been demonstrated in numerous organisms. We empirically demonstrate for Drosophila melanogaster that both sperm quality and sperm quantity independently contribute to competitive male fertilization success. In addition to these independent effects, there was a significant interaction between sperm quality and quantity that suggests an internal positive reinforcement on selection for sperm quality, with selection predicted to intensify as investment per sperm increases and the number of sperm competing declines. The mechanism underlying the sperm quality advantage is elucidated through examination of the relationship between female sperm-storage organ morphology and the differential organization of different length sperm within the organ. Our results exemplify that primary sex cells can bear secondary sexual straits.

Success in sperm competition, occurring whenever females mate with multiple males [1], is predicted to be influenced by variation in ejaculate quality and interactions among competing sperm [2]. Yet, apart from sperm number, relevant ejaculate characteristics and sperm-sperm interactions are poorly understood, particularly within a multivariate framework and the natural selective environment of the female reproductive tract. Here, we used isogenic lines of Drosophila melanogaster with distinguishable sperm to demonstrate and partition genetic variation in multiple sperm quality and performance traits. Next, by competing males from different lines, we show how rival sperm significantly influence each other's velocity and reveal that males with relatively slow and/or long sperm better displace rival sperm and resist displacement, thus avoiding ejection by the female from her reproductive tract. Finally, we establish fitness consequences of genetic variation in sperm quality and its role in securing a numerical advantage in storage by showing that offspring paternity is determined strictly by the representation of stored, competing sperm. These results provide novel insight into complex postcopulatory processes, illustrate that different ejaculate traits are critical at different biologically relevant time-points, and provide a critical foundation for elucidating the role of postcopulatory sexual selection in trait diversification and speciation.

In contrast to early predictions, it is now widely accepted that males incur substantive costs from ejaculate production. Hence, males are predicted to allocate their reproductive investments, including ejaculate size, relative to the risk of sperm competition and to female quality. The study of sperm allocation, however, has been technically challenging with nonvirgin females because sperm from different males must be discriminated within the female reproductive tract. To date, such investigations have thus largely been restricted to species that transfer sperm in spermatophores or for which females can be fitted with a harness to capture the incoming ejaculate. In this study, we examined sperm allocation using male Drosophila melanogaster that express a fluorescently labeled protein in sperm heads, allowing us to quantify sperm numbers from different males within the female reproductive tract. We found that male D. melanogaster deliver significantly more sperm to mated, large or young females compared with virgins, small or old females, respectively, whereas copulation duration was only significantly longer with large than with small females. These results provide further evidence for costly ejaculate production and consequent prudent allocation of sperm.

In some spiders, a discrete portion of the male's copulatory organ (the apical sclerite) breaks off during copulation and remains in the female's reproductive tract. Apical sclerites may prevent insemination by rivals (sperm competition), stimulate females to favourably bias paternity (cryptic choice) or breakage may reflect sexual conflict over copulation duration with little direct effect on paternity. It has been assumed that any benefits of organ breakage are balanced by a large cost (male sterility) in species where males could otherwise mate multiply, but this has never been experimentally tested. We examined these ideas in the Australian redback spider (Latrodectus hasselti Thorell 1870, Araneae: Theridiidae), a species where males are functionally sterile after one normal mating. We experimentally removed sclerites and found males were able to mate, had similar copulation durations and transferred similar numbers of sperm as males with intact sclerites. Benefits of o...

Many sperm competition studies have identified copulation duration as an important predictor of paternity. This result is often interpreted as a sperm transfer effect-it is assumed that sperm transfer is limited by copulation duration. Here we test the assumption of duration-dependent sperm transfer in the Australian redback spider, Latrodectus hasselti, in which a correlation between copulation duration and paternity has been implicated in the evolution of a rare male self-sacrifice behavior. Male redbacks facilitate sexual cannibalism by females during copulation. Sexual cannibalism is apparently adaptive for redback males, in part because it results in longer copulations (25 versus 11 min.), and copulation duration is positively correlated with paternity. We assessed sperm transfer in normal copulations and in copulations that we terminated at 5, 10, or 20 min. Our results show that the paternity advantage of sexual cannibalism is not owing to time-dependent sperm transfer, as redback males transfer the majority of their sperm within the first 5 min of copulation. This suggests that the link between copulation duration and paternity may instead be owing to cryptic female choice or the transfer of nongametic ejaculatory substances. Results further indicate that the act of cannibalism itself might play a role in mediating sperm transfer. This study highlights the importance of understanding mechanisms of sperm transfer when attempting to interpret the outcome of sperm competition studies.