High levels of multiple paternity in a spermcast mating freshwater mussel

Multiple paternity is an important characteristic of the genetic mating system and common across a wide range of taxa. Multiple paternity can increase within‐population genotypic diversity, allowing selection to act on a wider spectre of genotypes, and potentially increasing effective population size. While the genetic mating system has been studied in many species with active mating behavior, little is known about multiple paternity in sessile species releasing gametes into the water. In freshwater mussels, males release sperm into the water, while eggs are retained and fertilized inside the female (spermcast mating). Mature parasitic glochidia are released into the water and attach to the gills of fish where they are encapsulated until settling in the bottom substrate. We used 15 microsatellite markers to detect multiple paternity in a wild population of the freshwater pearl mussel (Margaritifera margaritifera). We found multiple paternity in all clutches for which more than two offspring were genotyped, and numbers of sires were extremely high. Thirty‐two sires had contributed to the largest clutch (43 offspring sampled). This study provides the first evidence of multiple paternity in the freshwater pearl mussel, a species that has experienced dramatic declines across Europe. Previous studies on other species of freshwater mussels have detected much lower numbers of sires. Multiple paternity in freshwater pearl mussels may be central for maintaining genetic variability in small and fragmented populations and for their potential to recover after habitat restoration and may also be important in the evolutionary arms race with their fish host with a much shorter generation time.     

Mate discrimination in Littorina littorea (L.) and L. saxatilis (Olivi) (Mollusca:Prosobranchia)

The ability of males of Littorina littorea and L. saxatilis to discriminate between mates of different sex, species and size was examined. In partner choice experiments males of L. littorea had the possibility to initiate a copulation with either a female or a male. The males did not show a preference for either sex. There was therefore no evidence that they could determine the sex of a conspecific prior to copulation. The duration of intrasexual copulation was considerably shorter than for intersexual copulation, both in the field and in laboratory experiments. For the two species, intersexual copulations were far more frequent than intrasexual ones. This can partly be explained by the difference in copulation time.Few interspecific copulating pairs were found on the shore. This may reflect a low interspecific encounter rate rather than a mechanism of species recognition. On all of these occasions, however, the active male was of L. saxatilis. It is argued that selection against precopulatory species and sex recognition is a more likely explanation than an hypothesis that states that the required mutations for precopulatory mate identification has not yet occurred. L. littorea males copulated longer with large than with small females. Copulation time was short with parasitized females, which are sterile or of low fecundity. The allocation of mating effort by males is discussed.     

Mating behaviour in Moina brachiata (Jurine, 1820) (Crustacea,Anomopoda)

Information on mating behaviour in Anomopoda isavailable for very few species only, though matelocation and recognition certainly play an importantrole in maintaining reproductive isolation betweenspecies. Ephippial females and males of Moinabrachiata were observed in a drop of water under amicroscope for 10–20 minutes. Different combinationsof specimens were used, but copulation was onlyobserved when two males and two ephippial females wereplaced together. Males were very active, and oftentried to grasp a female, whereas females usually madeattempts to escape during the entire period of mating.Three phases were recognized: capture, positioning andcopulation. The male captured the female from thedorsal side, then moved to the ventral side and tooka position with its length axis being perpendicular tothat of the female, forming a sort of cross.Thereafter the pair started rotating around the lengthaxis of the female, while the male pushed thepostabdomen into the females brood pouch. Copulationlasted from 16 to 25 seconds. When different kinds offemales were used, males showed preference forephippial females with an empty ephippium and enlargedovaries. Our results indicate that not only visual andtactile cues may be important in identifying speciesidentity and receptivity of the female, but alsochemical signals.     

Loading constraints,body size and mating preference in Gammarus species

Size relationships among pairing Gammarus were examined with reference to two hypotheses for sexual size dimorphism and assortative mating among aquatic Peracarida (Crustacea). The sizes of pairing and non-pairing animals were compared in different experimental conditions where the size of one or both sexes was controlled. The experimental results present a complex picture which suggests that both sexual selection and loading constraints are likely to play a role in determining mating decisions in these animals.Department of Adult & Continuing Education, University of Durham     

The energetic cost of mating in a promiscuous cephalopod

Costs that individuals incur through mating can play an important role in understanding the evolution of life histories and senescence, particularly in promiscuous species. Copulation costs, ranging from energy expenditure to reduced longevity, are widely studied in insects but have received substantially less attention in other taxa. One cost of mating, the energetic cost, is poorly studied across all taxa despite its potential importance for the many species where copulation is physically demanding and/or frequent. Here, we investigated the energetic cost of mating in both male and female dumpling squid (Euprymna tasmanica). In this species, copulation can last up to 3 h and requires that the male physically restrains the female. We report that the act of copulation halves the swimming endurance of both sexes, and that they take up to 30 min to recover. Such a reduction in post-copulatory performance may have important implications for predator avoidance, foraging ability and energy allocation. Therefore, quantifying this cost is essential to understand the evolution of reproductive strategies and behaviours such as female receptivity and male and female mating frequency.     

The mating behaviour of tsetse flies (Glossina): a review

Abstract. The mode of reproduction of tsetse flies, by adenotrophic viviparity, is unusual among the Diptera and is associated with many unique aspects of the tsetse's mating system. Tsetse exist at relatively low densities in the environment but a combination of olfactory and visual stimuli brings males and virgin females together on or around host animals. The behavioural repertoire associated with mate location and identification, courtship and copulation is regulated by external physical and chemical stimuli as well as by internal physiological mechanisms. With a view to identifying stimuli that could be used to manipulate tsetse behaviour and exploited for control purposes, much progress has been made in recent years in elucidating the mating behaviour of tsetse and its regulatory mechanisms. This progress and the current state of understanding of tsetse mating behaviour is reviewed.     

Analysis of the mating activity of male tsetse flies Glossina m. morsitans and G. pallidipes in the laboratory

ABSTRACT. The responses of male Glossina morsitans morsitans West-wood and Glossina pallidipes Austen to freeze-killed females were examined in the laboratory. Analyses were performed using a specially designed, automated, computer-based, recording system. G. pallidipes were more active than G. m. morsitans , interacting with the female decoys twice as often. Interactions with the decoys divided broadly into short-stay (<60 s) and long-stay, full copulatory attempts. For G. m. morsitans 90% of interactions resulted in full copulatory attempts, the mean duration of which was >1 h. For G. pallidipes only 40% of interactions resulted in full copulatory attempts, the mean duration of which was 35 min. The initiation of interactions showed a clear V-shaped activity pattern in G. m. morsitans but in G. pallidipes only a morning peak was observed. In neither species was there a tendency for full copulatory responses to be initiated in any specific period of the diurnal activity pattern. The results indicate that the two species have very different mating systems, and represent an initial step in the quantification of these differences.     

Nonrandom mating in pikas Ochotona princeps: evidence for inbreeding between individuals of intermediate relatedness

The extent and effect of inbreeding in natural populations remain largely undetermined. Pikas Ochotona princeps have been considered a likely candidate for close inbreeding in natural populations due to observations of frequent juvenile philopatry (colonization of natal home range or neighbouring home range) and high levels of spatial overlap and social tolerance between neighbouring individuals of the opposite sex. A 4-year investigation of inbreeding in pikas, however, has revealed that dispersal and mating patterns are uncoupled in this species, i.e. explained by different hypotheses. DNA fingerprinting analysis revealed that band-sharing scores between mated pairs, identified via parentage analysis, were not commensurate with band-sharing among known first-order relatives, but were similar to scores for a small sample of known second-order relatives (i.e. half-siblings, grandparent-grandchild pairs). Band-sharing scores between mated pairs were then compared with those between potential mated pairs within the population to assess whether mating was random or nonrandom with respect to genetic similarity. The results of Monte Carlo randomization tests show that pikas mated with individuals with intermediate genetic similarity in greater proportion than would be expected by chance. These data suggest mate choice in pikas may be based upon intermediate levels of relatedness.     

Factors influencing male mating behaviour in Gambusia affinis (Baird & Girard) with a coercive mating system

The effects of male and female body size, and correlated characteristics, on male mating behaviour were investigated in the western mosquitofish Gambusia affinis . Because larger females typically have larger broods in Gambusia sp., it was predicted that males would attempt more copulations with larger females. Two-way ANOVA showed that female body size was a significant predictor of male mating behaviour but male size was not. The effects of a suite of additional traits (both male and female) on male mating attempts were also tested. In a stepwise multiple regression, female standard length ( L _S), size of the female gravid spot and male testes mass were significant predictors of male mating attempts, accounting for c. 27% of variation in male mating. Path analysis showed that differences between male and female L _S, male body condition and male testes mass were significant predictors of male mating attempts, and also accounted for 27% of the variation in male mating attempts. The two statistical models were very similar in their predictive power, but differed slightly in significant predictor variables. Results confirm that factors other than female size are important predictors of male mating behaviour in the western mosquitofish.     

Variable Female Mating Positions and Offspring Sex Ratio in the Spider Pityohyphantes phrygianus (Araneae: Linyphiidae)

Chromosomal sex determination and male heterogamety have been thought to seriously impede direct sex ratio control. However, in Pityohyphantes phrygianus, a solitary sheetweb spider with a skewed sex ratio, earlier experimental studies suggested that there are options for female control of offspring sex ratio, if females change their position during the normal mating sequence. Here we show that under natural conditions there is considerable between-female variation in positions, especially after termination of mating. Computer simulations of the orientation of female inner genitalia suggest that sperm are placed in different storage sites depending on the positions adopted. This means that a specific position after mating might potentially influence offspring sex ratio. The variance in offspring sex ratio among females in earlier experiments was binomially distributed, which leads us to conclude that females control the mean sex ratio but do not exercise direct control of the sex of individual offspring.     

A field test of a model for the stability of androdioecy in the freshwater shrimp,Eulimnadia texana

The evolution of hermaphroditism from dioecy is a poorly studied transition. Androdioecy (the coexistence of males and hermaphrodites) has been suggested as an intermediate step in this evolutionary transition or could be a stable reproductive mode. Freshwater crustaceans in the genus Eulimnadia have reproduced via androdioecy for 24+ million years and thus are excellent organisms to test models of the stability of androdioecy. Two related models that allow for the stable maintenance of males and hermaphrodites rely on the counterbalancing of three life history parameters. We tested these models in the field over three field seasons and compared the results to previous laboratory estimates of these three parameters. Male and hermaphroditic ratios within years were not well predicted using either the simpler original model or a version of this model updated to account for differences between hermaphroditic types (‘monogenic’ and ‘amphigenic’ hermaphrodites). Using parameter estimates of the previous year to predict the next year's sex ratios revealed a much better fit to the original relative to the updated version of the model. Therefore, counter to expectations, accounting for differences between the two hermaphroditic types did not improve the fit of these models. At the moment, we lack strong evidence that the long‐term maintenance of androdioecy in these crustaceans is the result of a balancing of life history parameters; other factors, such as metapopulation dynamics or evolutionary constraints, may better explain the 24+ million year maintenance of androdioecy in clam shrimp.     

Intentional multiple mating by females in a species where sneak fertilization circumvents female choice for parental males

This paper describes how individual female ocellated wrasse Symphodus ocellatus distribute their spawning among males and nests in space and time. It is based on previously collected genetic data of larvae from ten different nests (used to reconstruct half and full‐sibling groupings both within and among nests on multiple days) and behavioural data of marked females across the reproductive season. Both the genetic analyses and behavioural observations confirm that female S. ocellatus intentionally engage in multiple mating, by repeatedly spawning at the same nest on different days and at several different nests (up to 12 spawning events over 3 weeks), leading to mixed paternity among her young. The main benefit of such high and intentional multiple mating is probably insurance against brood failure due to nest predation, desertion or poor paternal care by the male. These findings reveal that even in systems where females attempt to avoid male‐controlled mixed paternity, they may still engage in intentional multiple mating due to these potential benefits.     

Cost of mating in male Menochilus sexmaculatus (Fabricius) (Coleoptera: Coccinellidae)

Mating and reproduction not only provide benefits but can also incur substantial costs. An increasing number of recent studies have indicated that mate searching, mating, and ejaculate production requires large amount of energy which can result in high cost to males. Here, we examine the effect of number of matings of males and mating duration on mating and reproductive success in Menochilus sexmaculatus (Coleoptera: Coccinellidae) by subjecting them to 3 consecutive matings of 30, 60, 90 min, and complete mating (a) with no interval and (b) with interval of 24 hr between the matings. The results reveal a reduction in reproductive performance for both the setup and reduction is more when mating is consecutive without gap. This shows that between subsequent matings males require time to recuperate the sperm/ejaculate. As a male engages in a greater number of matings, the time it takes to begin mating when paired with a female increases and the overall duration of mating decreases. However, frequent mating attempts made by males result in reduction in their body weight. Thus, in this ladybird beetle mating and related physiological process consume lots of energy and the body weight of males tend to decrease which further modifies their reproductive performance. The study helps us to identify the cost of mating from male's perspective and indicates that males need to rejuvenate between the matings.     

The use of fluorescent pigments to study insect behaviour: investigating mating patterns in a butterfly population

Abstract. 1. Mating patterns in a natural population of checkerspot butterflies, Euphydryas editha Boisduval, were investigated by dusting male external genitalia with fluorescent dyes and examining mated females under UV light.
2. These dyes may have diverse applications in studies of insect populations.     

Effects of reproductive state and host resource experience on mating decisions in a walnut fly

Prior experience with conspecifics or essential resources, aswell as physiological condition, can have important influenceson an animal's reproductive behavior. While effects of experienceand physiological state (such as reproductive condition) aregenerally treated separately in theoretical discussions, theyoften interact. No previous study has attempted to distinguisheffects of experience on physiological state from other effectsof experience in the context of mating behavior. In a studyof a walnut-infesting tephritid fly (Rhagoletis juglandis),we examined the effects of host fruit experience on mating behavior.We manipulated physiological state in terms of egg load (definedas the number of mature oocytes in a female's ovaries) independentlyof fruit experience to distinguish the effects of these variables.We found that females with high egg loads were significantlymore likely to copulate than low–egg load females; thelevel of fruit experience had no effect on propensity to copulate,except via effects on egg load. In contrast, females with priorexposure to fruit copulated for a significantly shorter durationthan control females, while egg load had no effect on copulationduration. These results suggest that female reproductive conditionand exposure to essential resources can have important, albeitdiverse effects on mating behavior. We discuss how distinguishingdifferent types of variables may provide insight into sexualconflict over mating decisions, as well as which sex controlsspecific aspects of behavior.     

Local parasite pressures and host genotype modulate epigenetic diversity in a mixed‐mating fish

Parasite‐mediated selection is one of the main drivers of genetic variation in natural populations. The persistence of long‐term self‐fertilization, however, challenges the notion that low genetic variation and inbreeding compromise the host's ability to respond to pathogens. DNA methylation represents a potential mechanism for generating additional adaptive variation under low genetic diversity. We compared genetic diversity (microsatellites and AFLPs), variation in DNA methylation (MS‐AFLPs), and parasite loads in three populations of Kryptolebias hermaphroditus, a predomintanly self‐fertilizing fish, to analyze the potential adaptive value of DNA methylation in relation to genetic diversity and parasite loads. We found strong genetic population structuring, as well as differences in parasite loads and methylation levels among sampling sites and selfing lineages. Globally, the interaction between parasites and inbreeding with selfing lineages influenced DNA methylation, but parasites seemed more important in determining methylation levels at the local scale.     

Inter-sexual multimodal communication during mating in wild geladas: the leading role of females

Female primates can emit vocalizations associated with mating that can function as honest signals of fertility. Here, we investigated the role of mating calls and visual signals in female geladas (Theropithecus gelada). Because females have a central role in the gelada society and seem to solicit sexual interactions, we answered whether they emit vocalizations in conjunction with gazing to increase mating success probability. Before and during copulations, females can emit pre-copulation calls and copulation calls. For the first time, we identified a new female vocalization emitted at the final stage of copulations (end-copulation call), possibly marking the occurrence of ejaculation. We found that longer pre-copulation call sequences were followed by both prolonged copulations and the presence of end-copulation calls, thus suggesting that females use pre-copulation calls to ensure successful copula completion. Moreover, we found that different combinations of female vocal types and gazing had different effects on male vocal behavior and motivation to complete the copula. The analysis of the vocal and visual signals revealed a complex inter-sexual multimodal chattering with the leading role of females in the signal exchange. Such chattering, led by females, modulates male sexual arousal, thus increasing the probability of the copula success.