Polyandry contributes to Gonipterus platensis (Coleoptera: Curculionidae) rearing.

Background: Gonipterus platensis Marelli, 1926 (Coleoptera: Curculionidae) is one of the main defoliating beetles in Eucalyptus plantations. Biological control with egg parasitoids is one of the main control strategies for this pest and a constant supply of fresh host eggs is required to rear the parasitoids. Polyandry can influence Gonipterus oviposition by increasing female fecundity and fertility; however, the high density of individuals in laboratory colonies can lead to male harassment, resulting in lower reproduction rate. The aim of this study was to measure the effects of monoandry and polyandry on the reproduction of G. platensis and the effects of male harassment on laboratory rearing conditions. Methods: Reproductive parameters were compared between three treatments: monoandry, where the female was allowed to mate daily with the same male; no choice polyandry, where the female was allowed to mate daily with a different male; and polyandry with choice, where the female was allowed to mate daily, but with a choice between five different males. Another experiment varying the density of males was conducted to evaluate the effect of male harassment. Results: Polyandry with choice resulted in the longest period of oviposition, highest fecundity and highest number of eggs per egg capsules when compared to monoandrous females. No negative effect related to male harassment in the laboratory, such as decreased fertility, fecundity, or number of eggs per egg capsule, was detected. Conclusion: Polyandry contributes to mass rearing as it increases fecundity and oviposition period on females and there is no evidence of male harassment on G. platensis.

Does the potential strength of sexual selection differ between mating systems with and without defensive behaviours? A meta-analysis.

The Darwin-Bateman paradigm predicts that females enhance their fitness by being choosy and mating with high-quality males, while males should compete to mate with as many females as possible. In many species, males enhance their fitness by defending females and/or resources used by females. That is, males directly defend access to mating opportunities. However, paternity analyses have repeatedly shown that females in most species mate polyandrously, which contradicts traditional expectations that male defensive behaviours lead to monandry. Here, in an extensive meta-analysis, encompassing 109 species and 1026 effect sizes from across the animal kingdom, we tested if the occurrence of defensive behaviours modulates sexual selection on females and males. If so, we can illuminate the extent to which males really succeed in defending access to mating and fertilisation opportunities. We used four different indices of the opportunity for sexual selection that comprise pre-mating and/or post-mating episodes of selection. We found, for both sexes, that the occurrence of defensive behaviours does not modulate the potential strength of sexual selection. This implies that male defensive behaviours do not predict the true intensity of sexual selection. While the most extreme levels of sexual selection on males are in species with male defensive behaviours, which indicates that males do sometimes succeed in restricting females' re-mating ability (e.g. elephant seals, Mirounga leonina), estimates of the opportunity for sexual selection vary greatly across species, regardless of whether or not defensive behaviours occur. Indeed, widespread polyandry shows that females are usually not restricted by male defensive behaviours. In addition, our results indicate that post-mating episodes of selection, such as cryptic female choice and sperm competition, might be important factors modulating the opportunity for sexual selection. We discuss: (i) why male defensive behaviours fail to lower the opportunity for sexual selection among females or fail to elevate it for males; (ii) how post-mating events might influence sexual selection; and (iii) the role of females as active participants in sexual selection. We also highlight that inadequate data reporting in the literature prevented us from extracting effect sizes from many studies that had presumably collected the relevant data.

Are There Edge Effects on the Genetic Diversity of the Trap-Jaw Ant Odontomachus chelifer (Formicidae: Ponerinae) in a Neotropical Savanna Fragment? A First Assessment.

Habitat fragmentation is considered an important threat to biodiversity, increasing species exposure to edge effects. The Brazilian Cerrado savanna is considered a biodiversity hotspot and has been converted to small, isolated fragments due to human activities. Ant communities and colony survivorship are known to be affected by edge effects in Cerrado, but to date there is no information on the genetic diversity of ant colonies at the edge of fragmented areas. Here, we investigate if colony genetic diversity and structure of Odontomachus chelifer (Latreille) ants (Hymenoptera: Formicidae) are subject to edge effects in a Cerrado reserve in southeast Brazil. Using microsatellites, we evaluated the number of breeders (queens and males) and the genetic diversity in O. chelifer colonies located in the interior versus edge of a Cerrado fragment. All O. chelifer nests had multiple queens, which presented a low mating frequency. The number of breeders and most estimates of genetic diversity did not differ between colonies at the edge versus interior of the fragment. Genetic structure was not influenced by nest location as well. However, we detected a small and positive increase in the observed heterozygosity in colonies located at fragment edges. High heterozygosity is thought to be particularly important in fast-changing environments, such as edges, providing an advantage for genetic diversity. Further investigation is needed to assess in greater detail how habitat loss affects O. chelifer biology. Our study is a first step toward elucidating edge effects on genetic diversity of ant colonies, a topic still poorly explored in tropical environments.

First evidence of multiple paternity in the blue shark (Prionace glauca).

Multiple paternity (MP) is defined as the behaviour in which females successfully mate with multiple males leading to offspring from different sires within the same litter. MP seems to be frequent and an evolutionary advantage in elasmobranchs. Here the authors report for the first time the occurrence of MP in the cosmopolitan blue shark Prionace glauca L. The evidence, gathered via microsatellite genotyping of pregnant females and their embryos, suggests that MP is very frequent in this species. Knowledge of MP in P. glauca should help describe more precisely its reproductive biology and contribute to the management of its populations.

Photuris lugubris Female Fireflies Hunt Males of the Synchronous Firefly Photinus palaciosi (Coleoptera: Lampyridae).

Femmes fatales (Ff) are female fireflies that hunt and feed on the males of other firefly species that they attract by responding with glows or flashes to their bioluminescent signals. Here, we present field observations demonstrating that Photuris lugubris females are Ff of male Photinus palaciosi, a synchronous firefly exploited as a tourist attraction in the mountains of central Mexico. We show that the hunting success of the Ff is low, as observed in previous studies, suggesting that the impact of predation on the prey population is low. We present experimental data showing sex-specific hunting behaviour, since only female P. lugubris fed on P. palaciosi. We also present experimental data showing that at least some female P. lugubris mate multiple times; we discuss the implications of this discovery for the switch between the mating and hunting modes of Ff. We discuss open questions, as well as the possible impact of Ff on tourism focused on synchronous fireflies.

Regional variation in multiple paternity in the brown smooth-hound shark Mustelus henlei from the northeastern Pacific.

Multiple paternity (MP) in the brown smooth-hound shark (Mustelus henlei) was assessed in 15 litters (15 mothers and 97 embryos) collected in the northern Gulf of California of which 86.7% were sired by more than one male (i.e., from 2 to 4 sires). When taken together with results from previous studies, this record indicates that there is regional variation in MP in M. henlei in the northeastern Pacific. This pattern is associated with variations in the reproductive traits of each population (e.g., female size and litter size). In the Gulf of California, the results of a generalized linear model (GLZ) indicated that the litters of larger females had a higher probability of MP compared to those of smaller females.

Evidence of multiple paternity for the endemic Xingu River stingray.

Multiple paternity (MP) is a phenomenon observed for more than 30 elasmobranch species. The Batoidea is more specious than the Selachii, but only three studies of MP have been conducted on batoids. The occurrence of MP in freshwater stingrays was tested using microsatellite markers, which were developed for Potamotrygon leopoldi. Six mothers and their litters were genotyped, providing the first evidence of MP for Potamotrygonidae, with an MP frequency of 33%.

Queens remate despite traumatic mating in stingless bees.

Males can control female reproduction using genital plugs to impede access by rivals. In social bees, ants, and wasps, plugging may involve traumatic mating, with females being harmed. In stingless bees, chances are that plugs may promote ovarian activan, and are thought to ensure single mating-a general tendency among the social Hymenoptera. However, understanding on relationships between mating plugs, traumatic mating, and mating systems in stingless bees remains limited. To address this, we (1) compared mated queens of 7 Neotropical species to understand the patterns of copulatory marks in females and (2) compared pre- and post-mating genitalia of males and females in Melipona fasciculata to depict plug functional morphology. Data revealed an unprecedented consequence of mating in stingless bees: the characteristic marks left by mating plugs on female abdomens and the inferences that can be made from them. To our surprise, in 1 species M. fasciculata we found that queens retain the plug long after mating, and may carry it for the rest of their lives. All the other 6 species retained the plug for only a short period. Remated queens were only found in M. seminigra, whose multiple copulatory marks match previous findings of polyandry in this species. Our study shows that queens can remate, and suggests that male genital morphology may determine in part the time persistence of plugs. We conclude that traumatic mating plugs do not fully prevent remating in stingless bees and that mating systems are not uniform in this group. Nonetheless, exceptional cases of facultative polyandry in social insects-for example, when mating plugs fail-may confirm a general tendency for single mating in close link with efficient mating plugs.

The first reproductive parameters and evidence of multiple paternity in one new spiny dogfish species, Squalus albicaudus (Squaliformes, Squalidae).

Understanding elasmobranch reproductive biology is necessary for species conservation. Multiple paternity (MP) has been reported for elasmobranchs, and this study investigates the reproductive aspects and mating system (paternity genetic analyses) for Squalus albicaudus. Thirteen pregnant females were analysed concerning reproductive parameters, and the mating system was assessed for nine females and their litters. The study found a mean fecundity of 2.84 pups per litter without correlation between total female length and the number of embryos per litter. One litter showed evidence of MP, indicating the presence of polyandrous behaviour of the species.

An atypical mating system in a neotropical manakin.

Most of the diversity in the mating systems of birds and other animals comes at higher taxonomic levels, such as across orders. Although divergent selective pressures should lead to animal mating systems that diverge sharply from those of close relatives, opportunities to examine the importance of such processes are scarce. We addressed this issue using the Araripe manakin (Antilophia bokermanni), a species endemic to a forest enclave surrounded by xeric shrublands in Brazil. Most manakins exhibit polygynous lekking mating systems that lack territoriality but exhibit strong sexual selection. In sharp contrast, we found that male Araripe manakins defended exclusive territories, and females nested within male territories. However, territoriality and offspring paternity were dissociated: males sired only 7% of nestlings from the nests within their territories and non-territorial males sired numerous nestlings. Moreover, female polyandry was widespread, with most broods exhibiting mixed paternity. Apparently, territories in this species function differently from both lekking arenas and resource-based territories of socially monogamous species. The unexpected territoriality of Araripe manakins and its dissociation from paternity is a unique evolutionary development within the manakin clade. Collectively, our findings underscore how divergences in mating systems might evolve based on selective pressures from novel environmental contexts.

Promiscous female Euxesta eluta derive nutrients for ovarian development by expelling and consuming ejaculates.

Seminal gifts range from important material donations to items that provide little direct benefit to females. Promiscuous, female silk corn flies Euxesta eluta expel and consume male ejaculates immediately after mating. The evolution and function of this peculiar behavior are currently unknown. We performed a series of experiments aimed to: determine if females under different dietary regimes derive nutrients or water for survival and/or reproduction from ejaculate consumption, if males suffer a fitness cost from supplying females with ejaculates, and if females prefer to mate and/or are more likely to store sperm from well fed than nutritionally stressed presumably inferior males. Experiments revealed that protein deprived E. eluta females derive nutrients for ovarian development through consumption of ejaculates of protein fed males. No seminal products affecting survival appear to be transferred in the consumed ejaculate. However, ovarian development, in contrast to testes growth, occurs in detriment of longevity. Females preferred to mate with protein fed males, yet sperm retention in spermathecae was extremely rare after a single mating. This finding suggests that females could be exerting post copulatory control. A key question that remained to be addressed for the understanding of this puzzling and promiscuous mating system is what ecological factors or male traits drive females to retain sperm from one or several males in order to achieve and/or maximize fertilization potential.

Polyandry and trade-off between fecundity and longevity in female Dichelops furcatus (Hemiptera: Pentatomidae).

A trade-off is a relationship between two life history characteristics principally reproduction and adult longevity that are fundamental in predicting the optimal life history in any given environment. Mating is indispensable for sexual reproduction, but also can impose risks to females. Nevertheless, in the majority of insects, females allow multiple mating. Dichelops furcatus (Hemiptera: Pentatomidae) is a pest of wheat and corn in Argentina and southern Brazil, but little is known about its reproduction (i.e., the characteristics of the process that results in offspring). We analyzed reproductive attributes of D. furcatus, and the effect of single mating vs. multiple matings, evaluating the trade-off between fecundity and adult female longevity. We found that mating is not required for D. furcatus to oviposit, and multiple copulations were costly in terms of reduced longevity. Although multicopulated females lived a shorter period, only the pre- and post-reproductive periods were shortened. Fecundity was not affected but fertility was incremented in multicopulated females. Females copulated only once oviposited most of the eggs in the first half of the reproductive period, while eggs oviposited in the second half were all inviable (did not hatch). Studying demographic attributes of phytophagous insects provides relevant information to better understand the population dynamics of pests.

Adult sex ratio influences mate choice in Darwin's finches.

The adult sex ratio (ASR) is an important property of populations. Comparative phylogenetic analyses have shown that unequal sex ratios are associated with the frequency of changing mates, extrapair mating (EPM), mating system and parental care, sex-specific survival, and population dynamics. Comparative demographic analyses are needed to validate the inferences, and to identify the causes and consequences of sex ratio inequalities in changing environments. We tested expected consequences of biased sex ratios in two species of Darwin's finches in the Galápagos, where annual variation in rainfall, food supply, and survival is pronounced. Environmental perturbations cause sex ratios to become strongly male-biased, and when this happens, females have increased opportunities to choose high-quality males. The choice of a mate is influenced by early experience of parental morphology (sexual imprinting), and since morphological traits are highly heritable, mate choice is expressed as a positive correlation between mates. The expected assortative mating was demonstrated when the Geospiza scandens population was strongly male-biased, and not present in the contemporary Geospiza fortis population with an equal sex ratio. Initial effects of parental imprinting were subsequently overridden by other factors when females changed mates, some repeatedly. Females of both species were more frequently polyandrous in male-biased populations, and fledged more offspring by changing mates. The ASR ratio indirectly affected the frequency of EPM (and hybridization), but this did not lead to social mate choice. The study provides a strong demonstration of how mating patterns change when environmental fluctuations lead to altered sex ratios through differential mortality.

How Efficient Is Fertilization by Traumatic Insemination in Orius insidiosus (Hemiptera: Anthocoridae)?

Traumatic insemination (TI) can be injurious to females, and females have evolved various paragenital structures to mitigate these impacts. We examined the mating behavior of Orius insidiosus (Say) and the consequences of single and double matings for female fitness. A total of 100 virgin females (4-6-d old) were directly observed while they mated with virgin males. Some of these females were mated a second time with a different, nonvirgin male 3-5 d later, after they oviposited in sunflower stems. Females were held in isolation, fed eggs of Ephestia kuehniella Zeller, and reproductive success was tracked for 30 d. Six females died during their first copulation (6%), and another within 48 h, without laying eggs. Four percent of the females died during their second copulations. Copulations lasting less than 90 s usually did not result in successful fertilization, and duration of copula was positively correlated with egg fertility in singly-mated females. Duration of copula was more than halved in second matings, twice as variable, and negatively correlated with 30 d fecundity. Thirty-seven percent of singly-mated females and 31% of twice-mated females were infertile, with fewer than half of all females producing 88% of all eggs. We conclude that O. insidiosus females are likely monandrous in the wild, and that TI in this species is inefficient, contributing to high variation in female fitness. Thus, mating involves a significant mortality risk for females, despite their possession of complex paragenital structures that ostensibly mitigate copulatory injury.

Floral meristem size and organ number correlation in Eucryphia (Cunoniaceae).

We present a comparative flower ontogenetic study in five species of the genus Eucryphia with the aim of testing whether differences in the organ number observed can be explained by changes in the meristematic size of floral meristem and floral organs. Species native to Oceania, viz. E. milliganii, E. lucida and E. moorei, have the smallest gynoecia with ca. 6 carpels, while the Chilean E. glutinosa and E. cordifolia present more than ten carpels. E. milliganii has the smallest flower with the lowest stamen number (ca. 50), while the other species produce around 200 stamens and more. Standardized measurements of meristematic sectors were taken in 49 developing flowers that were classified into three well-defined ontogenetic stages. Sizes of meristems varied significantly among species within each developmental stage as revealed by ANOVA analyses. Significant regressions between organ number and corresponding meristem size were consistent with the premise that a larger meristem size prior to organ initiation could be determining for a higher organ number. Flower organogenesis in Eucryphia also involves relevant meristem expansion while the organs are initiated, which results in a particular androecium patterning with a chaotic stamen arrangement. Meristem expansion also appears to be slower but more extensive in species with larger initial meristematic size, suggesting that flower phenotype can be determined in ontogeny by this heterochronic interplay of space and time.

Females of a gift-giving spider do not trade sex for food gifts: a consequence of male deception?

BACKGROUND: Polyandry is commonly maintained by direct benefits in gift-giving species, so females may remate as an adaptive foraging strategy. However, the assumption of a direct benefit fades in mating systems where male gift-giving behaviour has evolved from offering nutritive to worthless (non-nutritive) items. In the spider Paratrechalea ornata, 70% of gifts in nature are worthless. We therefore predicted female receptivity to be independent of hunger in this species. We exposed poorly-fed and well-fed females to multiple males offering nutritive gifts and well-fed females to males offering worthless gifts. RESULTS: Though the treatments strongly affected fecundity, females of all groups had similar number of matings. This confirms that female receptivity is independent of their nutritional state, i.e. polyandry does not prevail as a foraging strategy. CONCLUSIONS: In the spider Pisaura mirabilis, in which the majority (62%) of gifts in nature are nutritive, female receptivity depends on hunger. We therefore propose that the dependence of female receptivity on hunger state may have evolved in species with predominantly nutritive gifts but is absent in species with predominantly worthless gifts.

Multiple Paternity in the Norway Rat, Rattus norvegicus, from Urban Slums in Salvador, Brazil.

The Norway rat, Rattus norvegicus, is one of the most important pest species globally and the main reservoir of leptospires causing human leptospirosis in the urban slums of tropical regions. Rodent control is a frequent strategy in those settings to prevent the disease but rapid growth from residual populations and immigration limit the long-term effectiveness of interventions. To characterize the breeding ecology of R. norvegicus and provide needed information for the level of genetic mixing, which can help identify inter-connected eradication units, we estimated the occurrence of multiple paternity, distances between mothers and sires, and inbreeding in rats from urban slum habitat in Salvador, Brazil. We genotyped 9 pregnant females, their 66 offspring, and 371 males at 16 microsatellite loci. Multiple paternity was observed in 22% (2/9) of the study litters. Of the 12 sires that contributed to the 9 litters, we identified 5 (42%) of those sires among our genotyped males. Related males were captured in close proximity to pregnant females (the mean inter-parent trapping distance per litter was 70 m, ±58 m SD). Levels of relatedness between mother-sire pairs were higher than expected and significantly higher than relatedness between all females and non-sire males. Our findings indicate multiple paternity is common, inbreeding is apparent, and that mother-sire dyads occur in close proximity within the study area. This information is relevant to improve the spatial definition of the eradication units that may enhance the effectiveness of rodent management programs aimed at preventing human leptospirosis. High levels of inbreeding may also be a sign that eradication efforts are successful.

Tamarins: Insights into monogamous and non-monogamous single female social and breeding systems.

Tamarins are reported to live in small multimale-multifemale groups characterized by a single breeding female. Here we present information on the composition and genetic relatedness of individuals in 12 wild-trapped groups of Weddell's saddleback tamarins (Saguinus weddelli) from northern Bolivia to determine if groups are best described as nuclear or extended families suggesting social monogamy or whether groups contain several unrelated same sex adults indicative of social polyandry/polygyny. Mean group size was 6.25 including an average of 2.16 adult males (range 1-4) and 2.08 adult females (1-3). No group contained only one adult male and one adult female and 25% of groups contained two parous females. We estimated the genetic relatedness among individuals using 13 polymorphic microsatellite markers. Across the population, mean relatedness was low and not significantly different among adult males versus among adult females, suggesting that both sexes disperse from their natal groups. Adults of both sexes also tended to have close same-sex adult relatives within their groups; relatedness among adult females of the same group averaged 0.31 and among adult males was 0.26. This suggests that tamarins of one or both sexes sometimes delay dispersal and remain as adults in their natal group or that emigration of same-sexed relatives into the same group may be common. Finally, parentage analyses indicated that, whereas the parents of juveniles generally were present in the group, this was not always the case. Based on these data, published reports of the presence of multiple breeding males and occasionally multiple breeding females in the same group, and the fact that less than 10% of groups in the wild contain a single adult male-adult female pair, we argue that social polyandry best characterizes the composition of tamarin groups and that monogamy is not a common mating pattern in Saguinus weddelli or other tamarin species.

Assessing paternity in japanese quails (Coturnix japonica) using microsatellite markers - inferences for its mating system and reproductive success

Microsatellite markers were analyzed in Japanese quails, Coturnix japonica, using different methodologies (PAGE and automated genotyping), in order to evaluate their use in paternity testing. Ten animal triplets composed by a female and two males were used to mate and generate an offspring. Paternity was determined in five-day-old embryos, and the data generated by fluorescent labeled and tailored primers in PCR and further automated genotyping were robust. Three microsatellite markers were polymorphic (Na = 5-8, H E = 0.75) and no loci were found to deviate significantly from Hardy-Weinberg equilibrium or showed any evidence of linkage disequilibrium (p > 0.05). A slight heterozygote deficiency and some incompatibilities between the female known parent and its offspring that involved homozygous genotypes were observed at GUJ0001 locus and may indicate the presence of null alleles. Although a reduced set of microsatellite primers were applied, it was possible to determine the paternity of 96.87% of the embryos, using combined data of three loci. The approach was useful for parentage inferring in a captive population of C. japonica and the results evidenced a potential polyandric mating system in the species, in which no advantage mechanism of last-male sperm precedence seems to occur.(AU)

Assessing paternity in japanese quails (Coturnix japonica) using microsatellite markers - inferences for its mating system and reproductive success

Microsatellite markers were analyzed in Japanese quails, Coturnix japonica, using different methodologies (PAGE and automated genotyping), in order to evaluate their use in paternity testing. Ten animal triplets composed by a female and two males were used to mate and generate an offspring. Paternity was determined in five-day-old embryos, and the data generated by fluorescent labeled and tailored primers in PCR and further automated genotyping were robust. Three microsatellite markers were polymorphic (Na = 5-8, H E = 0.75) and no loci were found to deviate significantly from Hardy-Weinberg equilibrium or showed any evidence of linkage disequilibrium (p > 0.05). A slight heterozygote deficiency and some incompatibilities between the female known parent and its offspring that involved homozygous genotypes were observed at GUJ0001 locus and may indicate the presence of null alleles. Although a reduced set of microsatellite primers were applied, it was possible to determine the paternity of 96.87% of the embryos, using combined data of three loci. The approach was useful for parentage inferring in a captive population of C. japonica and the results evidenced a potential polyandric mating system in the species, in which no advantage mechanism of last-male sperm precedence seems to occur.