Evolution of Size-Fecundity Relationship in Medaka Fish From Different Latitudes.

In most fishes, the number of offspring increases with maternal body size. Although this size-fecundity relationship often varies among species as a result of the coevolution of life-history traits, the genetic basis of such size-fecundity relationships remains unclear. We explored the genetic basis underlying this size-fecundity relationship in two small medaka species, Oryzias latipes and O. sakaizumii. Our findings showed that O. sakaizumii has a higher fecundity than O. latipes, and quantitative trait locus analysis using interspecific F2 hybrids showed that chromosome 23 is linked to the size-fecundity relationship. In particular, the genes igf1 and lep-b in this region are known to be associated with life-history traits, including somatic growth, gonad maturation, and progeny numbers in various taxa. Because O. sakaizumii is distributed at higher latitudes and has a shorter spawning season than O. latipes in the wild, we propose that the relatively high fecundity observed in O. sakaizumii is an adaptation to high latitudes. We also discuss the potential ecological ramifications associated with the evolution of increased fecundity in this species.

Density-dependent mating behaviors reduce male mating harassment in locusts.

Male mating harassment may occur when females and males do not have the same mating objectives. Communal animals need to manage the costs of male mating harassment. Here, we demonstrate how desert locusts in dense populations reduce such conflicts through behaviors. In transient populations (of solitarious morphology but gregarious behavior), we found that nongravid females occupied separate sites far from males and were not mating, whereas males aggregated on open ground (leks), waiting for gravid females to enter the lekking sites. Once a male mounted a gravid female, no other males attacked the pair; mating pairs were thereby protected during the vulnerable time of oviposition. In comparison, solitarious locusts displayed a balanced sex ratio in low-density populations, and females mated irrespective of their ovarian state. Our results indicate that the mating behaviors of desert locusts are density dependent and that sex-biased behavioral group separation may minimize the costs of male mating harassment and competition.

Bateman gradients reflect variation in sexual selection in a species with dynamic sex roles.

Bateman gradients, the slope of the regression of reproductive success on mating success, are among the most commonly reported measures of sexual selection. They are particularly insightful in species with reversed sex roles, where females are expected to be under sexual selection. We measured Bateman gradients in replicate experimental populations of the spermatophore gift-giving bushcricket Kawanaphila nartee (Orthoptera: Tettigoniidae). In this species, the operational sex ratio (OSR) and thus the sex competing for mates varies depending on the availability of pollen food resources: under pollen-limited regimens females are more competitive, whereas under pollen-rich regimens males are more competitive. We maintained populations in enclosures with either limited or supplemented pollen and calculated Bateman gradients for males and females under both conditions. Bateman gradients were significantly positive in males, and the slope was steeper in pollen-supplemented populations where the OSR was more male-biased. Bateman gradients for females were shallow and nonsignificant regardless of pollen availability. Our results show that the strength of sexual selection on males can depend on environmental context. The lack of significant gradients among females may reflect experimental limitations on our ability to estimate Bateman gradients in female K nartee.

Effects of operational sex ratio, mating age, and male mating history on mating and reproductive behavior in Grapholita molesta.

The short-lived polygamous moth Grapholita molesta (Busck) is an important fruit pest worldwide. Trapping males by synthetic female sex pheromones is not an effective reproductive control strategy. It is important to improve this technology by understanding the mating system of G. molesta. This study investigated mating opportunities and fertile egg production by altering the operational sex ratio, mating age, and male mating history in repeated single mating and multiple mating in the two sexes. Our results showed that the mating and reproductive parameters of virgin males were affected by the number and age of virgin females. Males preferred a female number ≤three-fifths of the male number or ≤2-day-old females, while they discriminated against a female number ≥three times of the male number or ≥5-day-old females. On the other hand, the mating and reproductive parameters of virgin females were affected by repeated single mating and especially multiple mating under different male mating histories. Females preferred once-mated males and discriminated against virgin males. These results indicated that mating systems including more and older virgin females for virgin males and different virgin males for virgin females may be suitable for suppressing G. molesta populations. Hence, these results revealed that preventing mating of virgin adults by synthetic female sex pheromones should be most effective in controlling G. molesta.

Relative Brain Size Is Predicted by the Intensity of Intrasexual Competition in Frogs.

Competition over mates is a powerful force shaping trait evolution. For instance, better cognitive abilities may be beneficial in male-male competition and thus be selected for by intrasexual selection. Alternatively, investment in physical attributes favoring male performance in competition for mates may lower the resources available for brain development, and more intense male mate competition would coincide with smaller brains. To date, only indirect evidence for such relationships exists, and most studies are heavily biased toward primates and other homoeothermic vertebrates. We tested the association between male brain size (relative to body size) and male-male competition across N=30 species of Chinese anurans. Three indicators of the intensity of male mate competition-operational sex ratio (OSR), spawning-site density, and male forelimb muscle mass-were positively associated with relative brain size, whereas the absolute spawning group size was not. The relationship with the OSR and male forelimb muscle mass was stronger for the male than for the female brains. Taken together, our findings suggest that the increased cognitive abilities of larger brains are beneficial in male-male competition. This study adds taxonomic breadth to the mounting evidence for a prominent role of sexual selection in vertebrate brain evolution.

Deer Indirectly Alter the Reproductive Strategy and Operational Sex Ratio of an Unpalatable Forest Perennial.

Environmental conditions impose restrictions and costs on reproduction. Multiple reproductive options exist when increased reproductive costs drive plant populations toward alternative reproductive strategies. Using 4 years of demographic data across a deer impact gradient, where deer alter the abiotic environment, we parameterize a size-dependent integral projection model for a sexually labile and unpalatable forest perennial to investigate the demographic processes driving differentiation in the operational sex ratio (OSR) of local populations. In addition to a relative increase in asexual reproduction, our results illustrate that nontrophic indirect effects by overabundant deer on this perennial result in delayed female sex expression to unsustainably large plant sizes and lead to more pronounced plant shrinkage following female sex expression, effectively increasing the cost of reproduction. Among plants of reproductive age, increased deer impact decreases the size-dependent probability of flowering and reduces reproductive consistency over time. This pattern in sex expression skews populations toward female-biased OSRs at low deer impact sites and male-biased OSRs at intermediate and high deer impact sites. While this shift toward a male-biased OSR may ameliorate pollen limitation, it also decreases the effective population size when coupled with increased asexual reproduction. The divergence of reproductive strategies and reduced lifetime fitness in response to indirect deer impacts illustrate the persistent long-term effects of overabundant herbivores on unpalatable understory perennials.

Male-only care and cuckoldry in black coucals: does parenting hamper sex life?

Providing parental care often reduces additional mating opportunities. Paternal care becomes easier to understand if trade-offs between mating and caring remain mild. The black coucal Centropus grillii combines male-only parental care with 50% of all broods containing young sired by another male. To understand how much caring for offspring reduces a male's chance to sire additional young in other males' nests, we matched the production of extra-pair young in each nest with the periods during which potential extra-pair sires were either caring for offspring themselves or when they had no own offspring to care for. We found that males which cared for a clutch were not fully excluded from the pool of competitors for siring young in other males' nests. Instead, the relative siring success showed a temporary dip. Males were approximately 17% less likely to sire young in other males' nests while they were incubating, about 48% less likely to do so while feeding nestlings, followed by 26% when feeding fledglings, compared to the success of males that currently did not care for offspring. These results suggest that real-life care situations by males may involve trade-off structures that differ from, and are less strict than those frequently employed in theoretical considerations of operational sex ratios, sex roles and parenting decisions.

Sperm morphology and count vary with fine-scale changes in local density in a wild lizard population.

Given that sperm production can be costly, theory predicts that males should optimally adjust the quantity and/or quality of their sperm in response to their social environment to maximize their paternity success. Although experiments demonstrate that males can alter their ejaculates in response to manipulations of the social environment and studies show that ejaculate traits covary with social environment across populations, it is unknown whether individual variation in sperm traits corresponds to natural variation found within wild populations. Using an island population of brown anole lizards (Anolis sagrei), we tested the prediction that sperm traits (sperm count, sperm morphology, sperm velocity) respond to natural variation in the risk of sperm competition, as inferred from the local density and operational sex ratio (OSR) of conspecifics. We found that males living in high-density areas of the island produced relatively larger sperm midpieces, smaller sperm heads, and lower sperm counts. Sperm traits were unrelated to OSR after accounting for the covariance between OSR and density. Our findings broaden the implications of sperm competition theory to intrapopulation social environment variation by showing that sperm count and sperm morphology vary with fine-scale differences in density within a single wild population.

Operational sex ratio predicts the opportunity and direction of sexual selection across animals.

The operational sex ratio (OSR) has long been assumed to be a key ecological factor determining the opportunity and direction of sexual selection. However, recent theoretical work has challenged this view, arguing that a biased OSR does not necessarily result in greater monopolisation of mates and therefore stronger sexual selection in the mate-limited sex. Hence, the role of the OSR for shaping animal mating systems remains a conundrum in sexual selection research. Here we took a meta-analytic approach to test whether OSR explains interspecific variation in sexual selection metrics across a broad range of animal taxa. Our results demonstrate that the OSR predicts the opportunity for sexual selection in males and the direction of sexual selection in terms of sex differences in both the opportunity for sexual selection and the Bateman gradient (i.e. the selection differential of mating success), as predicted by classic theory.

Multiple paternity in a viviparous toad with internal fertilisation.

Anurans are renowned for a high diversity of reproductive modes, but less than 1 % of species exhibit internal fertilisation followed by viviparity. In the live-bearing West African Nimba toad (Nimbaphrynoides occidentalis), females produce yolk-poor eggs and internally nourish their young after fertilisation. Birth of fully developed juveniles takes place after 9 months. In the present study, we used genetic markers (eight microsatellite loci) to assign the paternity of litters of 12 females comprising on average 9.7 juveniles. In 9 out of 12 families (75 %), a single sire was sufficient; in three families (25 %), more than one sire was necessary to explain the observed genotypes in each family. These findings are backed up with field observations of male resource defence (underground cavities in which mating takes place) as well as coercive mating attempts, suggesting that the observed moderate level of multiple paternity in a species without distinct sperm storage organs is governed by a balance of female mate choice and male reproductive strategies.

Reproductive skew drives patterns of sexual dimorphism in sponge-dwelling snapping shrimps.

Sexual dimorphism is typically a result of strong sexual selection on male traits used in male-male competition and subsequent female choice. However, in social species where reproduction is monopolized by one or a few individuals in a group, selection on secondary sexual characteristics may be strong in both sexes. Indeed, sexual dimorphism is reduced in many cooperatively breeding vertebrates and eusocial insects with totipotent workers, presumably because of increased selection on female traits. Here, we examined the relationship between sexual dimorphism and sociality in eight species of Synalpheus snapping shrimps that vary in social structure and degree of reproductive skew. In species where reproduction was shared more equitably, most members of both sexes were physiologically capable of breeding. However, in species where reproduction was monopolized by a single individual, a large proportion of females--but not males--were reproductively inactive, suggesting stronger reproductive suppression and conflict among females. Moreover, as skew increased across species, proportional size of the major chela--the primary antagonistic weapon in snapping shrimps--increased among females and sexual dimorphism in major chela size declined. Thus, as reproductive skew increases among Synalpheus, female-female competition over reproduction appears to increase, resulting in decreased sexual dimorphism in weapon size.

Transgenerational plasticity mitigates the impact of global warming to offspring sex ratios.

Global warming poses a threat to organisms with temperature-dependent sex determination because it can affect operational sex ratios. Using a multigenerational experiment with a marine fish, we provide the first evidence that parents developing from early life at elevated temperatures can adjust their offspring gender through nongenetic and nonbehavioural means. However, this adjustment was not possible when parents reproduced, but did not develop, at elevated temperatures. Complete restoration of the offspring sex ratio occurred when parents developed at 1.5 °C above the present-day average temperature for one generation. However, only partial improvement in the sex ratio occurred at 3.0 °C above average conditions, even after two generations, suggesting a limitation to transgenerational plasticity when developmental temperature is substantially increased. This study highlights the potential for transgenerational plasticity to ameliorate some impacts of climate change and that development from early life may be essential for expression of transgenerational plasticity in some traits.

Effect of Female-Biased Sex Ratios on Female Homosexual Behavior in Japanese Macaques: Evidence for the "Bisexual Preference Hypothesis".

We aimed to explain the frequent and prevalent female homosexual behavior in the context of female-biased operational sex ratios (OSR) and qualified sex ratios (Q) in a free-ranging group of Japanese macaques (Macaca fuscata) living at Arashiyama-Kyoto, Japan. Our data included the average availability of sexually mature males during females' putative fertile period (OSR), the ratio of sexually mature males to sexually mature females (Q), as well as heterosexual and female homosexual solicitations and consortships collected during 13 mating seasons from 136 females. Our results did not support the "heterosexual deprivation hypothesis," which holds that female homosexual behavior is attributable to a shortage of male mates. Likewise, our results did not support the "lack of opposite-sex sexual competitor hypothesis," which holds that females have more access to female mates when male sexual rivals are scarce. Of the 11 predictions tested, only one yielded statistically significant results: we found that higher ratios of availability of preferred female partners to preferred male partners were associated with female homosexual consortships rather than female heterosexual consortships. This result supported the "bisexual preference hypothesis," which holds that female homosexual behavior is attributable to female preference for certain female mates relative to certain male mates. We conclude that when a female targets another female as a mate, it is an active choice for a female sexual partner over available male alternatives, rather than a by-default situation that occurs because males are not available as sexual partners, or because females are better able to access female sexual partners due to a scarcity of male sexual competitors.

How choosy should I be? The relative searching time predicts evolution of choosiness under direct sexual selection.

Most theoretical research in sexual selection has focused on indirect selection. However, empirical studies have not strongly supported indirect selection. A well-established finding is that direct benefits and costs exert a strong influence on the evolution of mate choice. We present an analytical model in which unilateral mate choice evolves solely by direct sexual selection on choosiness. We show this is sufficient to generate the evolution of all possible levels of choosiness, because of the fundamental trade-off between mating rate and mating benefits. We further identify the relative searching time (RST, i.e. the proportion of lifetime devoted to searching for mates) as a predictor of the effect of any variable affecting the mating rate on the evolution of choosiness. We show that the RST: (i) allows one to make predictions about the evolution of choosiness across a wide variety of mating systems; (ii) encompasses all alternative variables proposed thus far to explain the evolution of choosiness by direct sexual selection; and (iii) can be empirically used to infer qualitative differences in choosiness.

Within-season variation in sexual selection in a fish with dynamic sex roles.

The strength of sexual selection may vary between species, among populations and within populations over time. While there is growing evidence that sexual selection may vary between years, less is known about variation in sexual selection within a season. Here, we investigate within-season variation in sexual selection in male two-spotted gobies (Gobiusculus flavescens). This marine fish experiences a seasonal change in the operational sex ratio from male- to female-biased, resulting in a dramatic decrease in male mating competition over the breeding season. We therefore expected stronger sexual selection on males early in the season. We sampled nests and nest-holding males early and late in the breeding season and used microsatellite markers to determine male mating and reproductive success. We first analysed sexual selection associated with the acquisition of nests by comparing nest-holding males to population samples. Among nest-holders, we calculated the potential strength of sexual selection and selection on phenotypic traits. We found remarkable within-season variation in sexual selection. Selection on male body size related to nest acquisition changed from positive to negative over the season. The opportunity for sexual selection among nest-holders was significantly greater early in the season rather than late in the season, partly due to more unmated males. Overall, our study documents a within-season change in sexual selection that corresponds with a predictable change in the operational sex ratio. We suggest that many species may experience within-season changes in sexual selection and that such dynamics are important for understanding how sexual selection operates in the wild.

Adult sex ratio variation: implications for breeding system evolution.

Adult sex ratio (ASR) exhibits immense variation in nature, although neither the causes nor the implications of this variation are fully understood. According to theory, the ASR is expected to influence sex roles and breeding systems, as the rarer sex in the population has more potential partners to mate with than the more common sex. Changes in mate choice, mating systems and parental care suggest that the ASR does influence breeding behaviour, although there is a need for more tests, especially experimental ones. In the context of breeding system evolution, the focus is currently on operational sex ratios (OSRs). We argue that the ASR plays a role of similar importance and urge researchers to study the ASR and the OSR side by side. Finally, we plead for a dynamic view of breeding system evolution with feedbacks between mating, parenting, OSR and ASR on both ecological and evolutionary time scales.

Mate competition and resource competition are inter-related in sexual selection.

Sexual selection can be affected by the competition for limited breeding resources and/or the competition for limited mates. Although there is ample evidence for each type of competition by itself, little is known about their relative importance and interaction. To address these questions, we established 48 experimental breeding populations of the two-spotted goby (Gobiusculus flavescens), a substrate-breeding fish with paternal care. In three experimental treatments, males were limited in the access to either nest sites or mates or were provided with both nests and mates in excess. We quantified male competition behaviour (agonistic and courtship), the opportunity for selection and selection on male body size. Limited access to nests and mates produced similar opportunities for selection, but only limited access to mates increased male competitive behaviours and caused positive selection on male body size. Selection on body size in the mate-limited treatment was due both to larger males being more likely to take up nests and to larger males being more likely to mate once they had a nest. These findings demonstrate that resource and mate limitation can differ in their effects on sexual selection. The results also reveal that resource and mating competition can be highly inter-related and not always separated in time, implying that methods to disentangle the two processes must be chosen with care. Future research should consider experimental and analytical approaches similar to those of the present study in attempts to elucidate the interaction of resource and mating competition in animals.

Sex ratio influences the motivational salience of facial attractiveness.

The sex ratio of the local population influences mating-related behaviours in many species. Recent experiments show that male-biased sex ratios increase the amount of financial resources men will invest in potential mates, suggesting that sex ratios influence allocation of mating effort in humans. To investigate this issue further, we tested for effects of cues to the sex ratio of the local population on the motivational salience of attractiveness in own-sex and opposite-sex faces. We did this using an effort-based key-press task, in which the motivational salience of facial attractiveness was assessed in samples of faces in which the ratio of male to female images was manipulated. The motivational salience of attractive opposite-sex, but not own-sex, faces was greater in the own-sex-biased (high competition for mates) than in the opposite-sex-biased (low competition for mates) condition. Moreover, this effect was not modulated by participant sex. These results present new evidence that sex ratio influences human mating-related behaviours. They also present the first evidence that the perceived sex ratio of the local population may modulate allocation of mating effort in women, as well as men.

Males follow females during fissioning of a group of northern muriquis.

Although well documented in matrilocal primate species, group fission is still a poorly known phenomenon among patrilocal primates. In this paper we describe in detail a group fission event in the population of northern muriquis at the Reserva Particular do Patrimônio Natural-Feliciano Miguel Abdala in Caratinga, Minas Gerais, Brazil, using Social Network Analyses (SNA). Data on association patterns were collected during systematic observations from May 2002 to September 2005, and analyzed for dry (from May to October) and rainy seasons (from November to April). The fission process started with subgroup formation in the rainy season 2002-2003, and was completed by the dry season of 2003. By the dry season 2003, the parent group (Jaó) had fissioned to form a second mixed-sex group (Nadir) while a subgroup of males (MU) moved between the parent group and the newly established group. Before the Jaó group fission started (dry season 2002) and during its initial phases (rainy season 2002-2003), females that ultimately composed the daughter group (Nadir) were the most peripheral in the association network. In the rainy season 2002-2003, the median monthly (N=6) operational sex ratio (OSR) of Jaó group was 2.81. However, once Jaó females initiated the fissioning process to establish the Nadir group, the OSR was more favorable to males in the Nadir group than in the Jaó group. Our results suggest that males followed the females to escape an unfavorable OSR in their natal group.

Phenology and Breeding Mechanisms of Calamus lobbianus Becc. and Calamus pygmaeus Becc.

The floral biology of Calamus is relatively unknown except for several species. In this study, Calamus lobbianus and Calamus pygmaeus were selected to represent the non-climbing rattan of the Sundaland's flagellate group. Observations on phenology, floral rewards and floral visitors as well as experiments on the breeding mechanism and operational sex ratio were performed for both species. For both species, anthesis started in the early morning for pistillate and staminate plants, ended by late noon (staminate plants) but lasted till the next day in pistillate plants for both species. Although both species appeared to be aseasonal in flowering, C. lobbianus exhibited a male-biased population while C. pygmaeus did not exhibit any sex bias. Nectar was observed to be extruded from the base of the petals of C. lobbianus of the staminate flowers and sterile staminate flowers of the pistillate plants. The concentration and volume of the nectar of the staminate flower and sterile staminate flower of C. lobbianus peaked at c. 11% and 9 µL around 1100 (Day 1) and c. 13% and 8 µL around 0930 (Day 1), respectively, but only appeared as a layer of glistening exudate in C. pygmaeus. Floral scent was not detected in any of the inflorescences of both species. Several insect visitors were observed to be the primary visitors of both Calamus species which include two species of Tetragonula, a species of Liostenogaster sp., and Stenodyneriellus sp. Experiments on breeding mechanism of C. lobbianus and C. pygmaeus indicate that both species are most likely apomictic. C. pygmaeus is capable of vegetative propagation through the rooting at the tip of inflorescence.

Abstrak: Biologi flora Calamus kurang difahami kecuali untuk beberapa spesies. Dalam kajian ini, Calamus lobbianus dan Calamus pygmaeus telah dipilih untuk mewakili rotan tidak memanjat dalam kumpulan berflagelum di Sundaland. Pemerhatian fenologi, ganjaran bunga dan pelawat bunga serta eksperimen mekanisma pembiakan dan nisbah jantina beroperasi telah dijalankan bagi dua spesies tersebut. Antesis bagi pokok betina dan jantan bermula dari awal pagi dan berakhir lewat petang (pokok jantan) manakala antesis bagi pokok pistilate berlangsung sehingga hari seterusnya. Pemerhatian pembungaan bagi kedua-dua spesies adalah tidak bermusim tetapi pembungaan C. lobbianus mempamerkan bias ke populasi jantan manakala C. pygmaeus tidak mempamerkan bias terhadap kedua-dua jantina. Nektar hanya dihasilkan pada bahagian dasar kelopak bunga jantan dan bunga jantan steril diad di sebelah bunga betina C. lobbianus. Pengeluaran nektar bunga jantan C.lobbianus memuncak sekitar jam 1100 (hari pertama) dengan kepekatan 11% dan isipadu sebanyak 9 µL. Pengeluaran bunga jantan steril memuncak sekitar jam 0930 (haripertama) dengan kepekatan 13% dan isi padu sebanyak 8 µL manakala C. pygmaeushanya menghasilkan lapisan cecair berkilat. Bau bunga kedua-dua spesies tidak dapatdikesan pada jambak bunga. Dua spesies Tetragonula dan satu spesies Liostenogastersp. diperhatikan sebagai pelawat utama untuk kedua-dua spesis Calamus. Eksperimenterhadap mekanisma pembiakan menunjukan bagi kedua-dua spesies Calamus adalahberkemungkinan apomik. C. pygmaeus mempunyai keuapayaan untuk menjalankanpropagasi secara vegetatif melalui penakaran pada hujung jambak bunga.