Interpopulational variation of the mating system in the peacock blenny Salaria pavo

The ecology and mating system of two populations of the peacock blenny Salaria pavo that have been reported to differ in sexual behaviour were studied. In the Gulf of Trieste, a northern Adriatic Sea population lives in rocky shores, whilst in the Ria Formosa, an Atlantic population inhabits a coastal lagoon with sandy barrier islands. In the Gulf of Trieste, nest availability was found to be higher and nest aggregation lower than in Ria Formosa. Nesting males courted females more and in higher proportion, and inversely, females directed less courtship displays towards nest holders in the Gulf of Trieste than in Ria Formosa. The relative frequency of small female-like males that reproduce by parasitically fertilizing eggs in the larger males’ nests was lower in the Gulf of Trieste population. This variation in the sexual behaviour of nesting males and females is likely to reflect a plastic behavioural response to the varying ecological conditions. The difference between populations in the relative frequency of parasitic males suggests that the male alternative reproductive tactics are condition-dependent.     

Effects of androgens on social behavior and morphology of alternative reproductive males of the Azorean rock-pool blenny

In the Azorean rock-pool blenny Parablennius sanguinolentus parvicornis two sequential reproductive tactics occur. Larger and older males establish breeding territories, while some of the smaller males become attached to nest-holder territories, acting as satellites on these territories, which they help to defend while trying parasitic fertilizations when females go in the nests to spawn. In the present paper we tested the effects of the androgens 17alpha-methyltestosterone (MT) and 11-ketotestosterone (KT) in the expression of male secondary sex characters and bourgeois behavior in satellite males. One week after satellites were implanted with Silastic tubes containing MT, KT, or castor oil (control), androgen-treated satellites had developed male secondary sex traits such as longer and wider male-type genital papilla and anal glands that secrete a sex pheromone, both traits being less expressed or absent, respectively, in satellite males. Androgen treatment had no effect on the gonadosomatic index or on the development of the testicular gland. KT treatment had a positive effect on relative liver weight. In terms of behavior, androgen-implanted individuals were less aggressive both in a mirror test and toward females when these were introduced into their tanks. MT-treated individuals spend more time inside the provided nests. Only androgen-implanted satellites managed to have the females entering their nests. When given a chance in a group tank either to try to attract females to their own nest or to act as satellites of an already established nest-holder's nest, MT-implanted males spent significantly more time in their own nest than near the nest-holder nest. These data suggest that androgens, particularly testosterone, may be involved in mating tactic switching in this species.     

Male reproductive success and its behavioural correlates in a polygynous mammal,the Galápagos sea lion (Zalophus wollebaeki)

Sexual selection theory predicts competitive males and choosy females. Nevertheless, since molecular marker‐based studies, paternity outside the expected mating patterns has increasingly been described. Even in highly polygynous systems, where paternity is expected to be strongly skewed towards large, dominant males, alternative mating tactics have been suggested. We examined reproductive success in the polygynous Galápagos sea lion (Zalophus wollebaeki). Semiaquatic territoriality allows females to move freely and may lower the degree of polygyny otherwise suggested by both territorial behaviour and strong sexual dimorphism. We assigned paternities with 22 microsatellites and analysed how male reproductive success was related to size, dominance status, intra‐sexual agonistic behaviour, proximity to females, and attendance in the colony. Male behaviour was consistent across two seasons for all parameters under consideration. Attendance was by far the most important determinant of paternal success. Skew in reproductive success towards large, dominant males was weak and dominance status played no role. This appears to be caused by an extremely long reproductive season lasting five or more months, making it difficult for any male to monopolize receptive females. Females seem to choose displaying males that were present in the colony for a long time rather than dominance per se. Sexual dimorphism in Galápagos sea lions may thus be more influenced by selection for fasting than fighting ability. Our data provide further evidence for alternative mating tactics, as several males gained relatively high reproductive success despite short attendance and hardly any involvement in agonistic interactions.     

Effect of Season and Age on the Gonadosomatic Index of Milne-Edwards' Sifakas (Propithecus diadema edwardsi) in Ranomafana National Park,Madagascar

Aiming to explain individual variation in gonadosomatic index (testicular volume per body weight), we examined the relationship between gonadosomatic index (GI), age, and season in a wild, seasonally breeding lemur—the Milne-Edwards' sifaka (Propithecus diadema edwardsi). Between 1987 and 2000, we captured and measured male lemurs from Ranomafana National Park in Madagascar 69 times. We used linear and polynomial regressions and residuals to examine the effect of season on GI and the effect of age on GI, after accounting for the effect caused by season among males of breeding age. As the breeding season approached, GI increased significantly, and all individuals measured in both the breeding and nonbreeding season had larger GI values in the breeding season. Among males of breeding age, we found no relationship between age and GI, after accounting for seasonal effects. Age, season and body weight fail to explain the variation remaining in GI. We discuss sperm competition, breeding-group composition, and the presence of parents as additional sources of variation in GI.     

Size Dependent Male Reproductive Tactic in the Two-Spotted Goby (Gobiusculus flavescens)

Male investment in testes and sperm duct gland in the polygamous nest breeding two-spotted goby Gobiusculus flavescens (Fabricius) was investigated in relation to time in reproductive season and individual physical parameters. This small teleost fish is most likely the most abundant species found along the rocky shores of the North East Atlantic. The two-spotted goby has a single reproductive season, during which nest-caring males can raise several clutches of offspring. According to the literature the males are on average larger than the females. Here we report for the first time a population showing a reversal of this trend, with males on average being smaller than females, a difference likely caused by a large proportion of small males. Early in the breeding season these small males have typical sneaker characters, with relatively large testes and small seminal duct glands compared to the larger dominant territorial males. The presence of these two alternative male reproductive tactics is confirmed by histological studies, which shows the presence of sperm in the sperm duct glands (SDG) of smaller males, but not in the SDG of intermediate and larger males. To our knowledge, males with typical sneaker characters have not been reported in earlier studied populations of two-spotted goby. Interestingly we found that testes investment declined significantly over the course of the breeding season, and that this reduction was significantly more pronounced in small compared to the large males. Further, a significant increase in seminal duct gland (SDG) mass was observed for the smaller males over the breeding season. We propose that this indicates a possible shift in mating tactic by smaller males from a parasitic to a nest-holding tactic over the course of the breeding season. Thus, the observed size dependent plasticity in investment in SDG over time suggests that the reproductive tactic of G. flavescens is conditional, and possibly influenced by mate availability and male—male competition.     

Diversity of sexual dimorphism in electrocommunication signals and its androgen regulation in a genus of electric fish, Apteronotus

Gymnotiform electric fish emit an electric organ discharge that, in several species, is sexually dimorphic and functions in gender recognition. In addition, some species produce frequency modulations of the electric organ discharge, known as chirps, that are displayed during aggression and courtship. We report that two congeneric species (Apteronotus leptorhynchus and A. albifrons) differ in the expression of sexual dimorphism in these signals. In A. leptorhynchus, males chirp more than females, but in A. albifrons chirping is monomorphic. The gonadosomatic index and plasma levels of 11-ketotestosterone were equivalent in both species, suggesting that they were in similar reproductive condition. Corresponding to this difference in dimorphism, A. leptorhynchus increases chirping in response to androgens, but chirping in A. albifrons is insensitive to implants of testosterone, dihydrotestosterone or 11-ketotestosterone. Species also differ in the sexual dimorphism and androgen sensitivity of electric organ discharge frequency. In A. leptorhynchus, males discharge at higher frequencies than females, and androgens increase electric organ discharge frequency. In A.␣albifrons, males discharge at lower frequencies than females, and androgens decrease electric organ discharge frequency. Thus, in both chirping and electric organ discharge frequency, evolutionary changes in the presence or direction of sexual dimorphism have been accompanied and perhaps caused by changes in the androgen regulation of the electric organ discharge. Accepted: 18 February 1998     

Alternative reproductive tactics are associated with sperm performance in invasive round goby from two different salinity environments

During male–male competition, evolution can favor alternative reproductive tactics. This often results in a dominant morph that holds a resource, such as a nest for egg laying, which competes with a smaller sneaker morph that reproduces by stealing fertilizations. The salinity environment can influence male growth rates, for example, via osmoregulatory costs, which in turn may influence the use of sneaker tactics for small males competing for mating opportunities. Salinity can also affect sperm directly; however, little is known of how salinity influences sneaker tactics through sperm performance. We sampled males of the invasive round goby (Neogobius melanostomus) from two environments, a freshwater river and a brackish estuary. This fish has two male morphs: nest‐holding dark males and non‐nest‐holding light males. We examined the role of water salinity of 0, 8, and 16 on sperm performance and found that for estuarine males, a salinity of 0 reduced sperm velocity compared to a salinity of 8 and 16. Riverine males had low velocity in all salinities. Sperm viability also decreased by over 30% in 0 salinity, compared to 8 and 16, for fish from both environments. Gobies produce ejaculate contents in specialized glands that could in theory shield sperm in an adverse environment. However, gland contents did not improve sperm performance in our tests. Body mass and age estimates indicate that riverine males invested more in somatic growth compared to estuarine males. Estuarine light morph males had a high enough gonadosomatic index to indicate sneaker tactics. We propose that when sperm performance is low, such as for the riverine males, sneaker tactics are ineffective and will be selected against or phenotypically suppressed. Instead, we interpret the increased investment in somatic growth found in riverine males as a life‐history decision that is advantageous when defending a nest in the next reproductive season.     

Age and reproductive investment in grass goby females in the Venice lagoon

The present work describes the relationship between age and reproductive investment in the grass goby females in the Venice Lagoon. Age was estimated by otoliths reading, while reproductive investment was assessed by either the relative number of mature females across the breeding season and their gonadosomatic index. Females from different size/age classes differed in the timing and level of reproductive investment. The oldest females (3+ years old) appeared earlier in the spawning habitat, investing more at the beginning of the breeding season. In turn a later arrival in the spawning habitats has been observed for younger females (0+) and smaller individuals of 1+ and 2+ age classes. Present results gave evidence of an earlier age/size at sexual maturity of the grass goby in the Venice Lagoon compared to previous information on the species from the same lagoon and from other Mediterranean areas. Results were discussed in the light of previous data on reproductive strategy of nesting males and of the relationship between fishing pressure and life history traits in the Venice lagoon.     

Gonad development and sperm motility of the diving beetle Cybister brevis Aubé, 1838 (Coleoptera: Dytiscidae) in response to seasonal changes in Japan

Gonad maturation in wild Cybister brevis Aubé, 1838 from Japan was investigated to determine the physiological mating season. Breeding experiments showed that hatching occurred from May to September, and the peak was observed in July. The gonadosomatic index (GSI) in collected females (ovary development) showed the highest value in May. The GSIs were low in other seasons. On the other hand, the GSIs in collected males (testes and accessory glands) remained at constant values during the experimental period. Interestingly, high sperm motility was exhibited in May and September, whereas it was low in December and March. This indicates the maturation difference between females and males. Females mature only at the beginning of the breeding season (May), but males do so from May to September. This result suggests that males have a wide-ranging maturation period and earlier sexual maturation than females, ensuring a successful fertilization.     

Sexual dimorphism in weight among the primates: The relative impact of allometry and sexual selection

In this paper, we examine allometric and sexual-selection explanations for interspecific differences in the amount of sexual dimorphism among 60 primate species. Based on evidence provided by statistical analyses, we reject Leutenegger and Cheverud’s [(1982). Int. J. Primatol.3:387-402] claim that body size alone is the major factor in the evolution of sexual dimorphism. The alternative proposed here is that sexual selection due to differences in the reproductive potential of males and females is the primary cause of sexual dimorphism. In addition, we propose that the overall size of a species determines whether the dimorphism will be expressed as size dimorphism,rather than in some other form.     

Brain and gonadal aromatase activity and steroid hormone levels in female and polymorphic males of the peacock blenny Salaria pavo

In the peacock blenny Salaria pavo large males with well-developed secondary sexual characters establish nests and attract females while small “sneaker” males mimic female sexual displays in order to approach the nests of larger males and parasitically fertilize eggs. These alternative reproductive tactics are sequential, as sneakers irreversibly switch into nesting males. This transition involves major morphologic and behavioral changes and is likely to be mediated by hormones. This study focuses on the role of aromatase, an enzyme that catalyses the conversion of androgens into estrogens, in the regulation of male sexual polymorphism in S. pavo. For this, sex steroid plasma levels and aromatase activity (AA) in gonads, whole brain and brain macroareas were determined in sneakers, transitional males (i.e. sneakers undergoing the transition into nesting males), nesting males and females collected in the field. AA was much higher in ovarian tissue than in testicular tissue and accordingly circulating estradiol levels were highest in females. This supports the view that elevated AA and estradiol levels are associated with the development of a functional ovary. Transitional males are in a non-reproductive phase and had underdeveloped testes when compared with sneakers and nesting males. Testicular AA was approximately 10 times higher in transitional males when compared with sneakers and nesting males, suggesting high AA has a suppressive effect on testicular development. Nesting males had significantly higher plasma levels of both testosterone (T) and 11-ketotestosterone when compared with the other male morphs and previous studies demonstrated that these androgens suppress female-like displays in sneakers. In the brain, AA was highest in macroareas presumably containing hypothalamic nuclei traditionally associated with the regulation of reproductive behaviors. Overall, females presented the highest levels of brain AA. In male morphs AA increased from sneakers, to transitional males, to nesting males in all brain macroareas. These results suggest that the transition into the nesting male tactic is accompanied both by an increase in testicular androgen production and by a higher conversion of androgens into estrogens in the brain. The increase in androgen production is likely to mediate the development of male secondary sexual characters while the increase in brain AA may be related to the behavioral changes associated with tactic transition.     

Differential investment into testes and sperm production in alternative male reproductive tactics of the African striped mouse (Rhabdomys pumilio)

Males that follow alternative reproductive tactics might differ in their investment into testis development and sperm production. The resource-allocation hypothesis predicts that males following a sneaker tactic should invest more into sperm production than dominant territorial males which should invest more into mate guarding. This hypothesis is supported by studies in species where individual males cannot switch between tactics (fixed tactics). Here we present the first data for a species where males can switch between tactics (plastic tactics). We studied African striped mice (Rhabdomys pumilio) in captivity, mimicking three tactics observed in the field: philopatric group-living males, singly-housed males representing roaming males, and group-living breeding males. We measured quantitative and qualitative reproductive traits, as well as serum and testis hormone concentrations. We found no support for the resource-allocation hypothesis, since breeding and singly-housed males invested similarly in testes and sperm. However, philopatric males had significantly smaller testes and epididymides, lower sperm counts, lower testosterone and higher corticosterone levels than males of the two other tactics. Philopatric males did not reach a larger body mass than singly-housed males with well developed reproductive traits, indicating that they did not trade investment in sperm production against growth. Interestingly, testis testosterone concentrations of philopatric males did not differ from those of other males. Our data suggest that philopatric males are reproductively suppressed by the breeding male, but might be ready to increase their serum testosterone levels when social and environmental conditions allow for this physiological switch accompanying the behavioral switch between tactics.     

Male sexual polymorphism, alternative reproductive tactics, and androgens in combtooth blennies (pisces: blenniidae)

In species in which intense intermale competition for the access to females is present males of lower competitive ability may adopt alternative reproductive tactics (ART) to get access to mates. These ART translate in many cases into male sexual polymorphism, with individuals following distinctly different tactics. Usually two alternative male morphs can be recognized in species with ART: (1) bourgeois males that compete for access to mates invest in typically male behaviors, such as building elaborated nests or displaying ornaments; and (2) parasitic males that take advantage of the success of the bourgeois males in attracting females and attempt "sneaker" fertilizations (e.g., sneaker and satellite males). In combtooth blennies (Blenniidae) the co-occurrence of ART and male sexual polymorphism has been described for two temperate species: the peacock blenny, Salaria pavo, and the Azorean rock-pool blenny, Parablennius sanguinolentus parvicornis. Interestingly, while in the peacock blenny the alternative male morph adopts a sneaker tactic, in the rock-pool blenny parasitic males act as satellites to nest-holder males. Thus, this variation in the ART expressed in these two closely related species allows for a comparative study of the proximate and ultimate factors affecting the expression of the two ART. In this article we summarize the available information on androgen levels in bourgeois and parasitic males of natural populations of the two species and of recent studies on the effect of exogenous administration of androgens on tactic switching in parasitic males of the two species. The information is discussed within the frame of the relative plasticity hypothesis, which predicts that plastic alternative morphs should show differences in hormone levels and that the administration of sex steroids should be effective in promoting the switch from the parasitic to bourgeois tactic. The evidence is only partly consistent with this hypothesis. Alternatively, a social transduction hypothesis that better fits the available data on androgens and ART in teleost is proposed. It states that the observed differences in androgen levels between alternative morphs should not be interpreted as an organization vs activation effect of steroids, but rather as the limited vs lifelong responsiveness of the neuroendocrine axis to social regulation.     

Reproductive strategy of Psectrogaster rhomboides Eigenmann & Eigenmann, 1889, a freshwater fish from Northeastern Brazil

Several environmental parameters related to the spawning season of the freshwater fish, Psectrogaster rhomboides were identified. The hypothesis predicts synchrony of the spawning season with the rainy period, presenting a seasonal reproductive strategy. This study investigated the total body length and body mass, sex ratio, body size at first sexual maturity, gonadosomatic index, condition factor, fecundity and reproductive period of P. rhomboides in the natural environment. Twelve monthly samplings (mean n = 20) were carried out from March 2009 to February 2010. Fish were captured using 4 cm diameter cast nets and environmental parameters such as rainfall, temperature, dissolved oxygen concentration, pH and electrical conductivity of the water were registered. The sampled population of P. rhomboides (n = 444) showed a sex ratio of 1 : 1. Females attained first sexual maturity at 15 cm total length and males at 15.3 cm total length. Spawning occurred from March to May, with high gonadosomatic index values during the rainy season. This species is a total spawner, with a mean fecundity of 6188 (±367) mature oocytes. P. rhomboides presents a seasonal reproductive strategy, providing a population increase and exploitation of the environmental resources during the rainy period. Rainfall (196.5 ± 4.9 mm) and the concentration of dissolved oxygen (7.82 ± 0.7018 mg L^?1) were influential factors during the spawning season. Gonadosomatic index and the condition factor were negatively correlated during the gonadal development cycle of this species. The monthly gonadosomatic index values show that spawning begins when the rainy season is at its maximum and ends synchronously (females r = 0.9171; males r = 0.889) with the end of the rainy season.     

Seasonal colour and antipredator behaviour in Etheostoma (Percidae)

This study examined how colour varies across season and sex in the fantail darter Etheostoma flabellare and the banded darter Etheostoma zonale. Etheostoma flabellare has male‐only parental care and exhibited slight sexual dimorphism in overall colour, with no discernible effect of season on colour; whereas E. zonale does not have parental care and exhibited substantial sexual dimorphism in colour, but only in the breeding season. Additionally, antipredator behaviour of E. zonale was compared between males that were fully coloured during the breeding season and males that were partially coloured at that time, but the effects of colour and season were not consistent across males.     

Sex,androgens, and whole‐organism performance in an Australian lizard

Understanding underlying physiological differences between the sexes in circulating androgens and how hormonal variation affects morphology–performance relationships may help clarify the evolution of sexual dimorphism in diverse taxa. Using a widely distributed Australian lizard (Eulamprus quoyii) with weak sexual dimorphism and no dichromatism, we tested whether circulating androgens differed between the sexes and whether they covaried with morphological and performance traits (bite force, sprint speed, endurance). Males had larger head dimensions, stronger bite force, faster sprint speed, and longer endurance compared to females. We found that the sexes did not differ in androgen concentrations and that androgens were weakly associated with both morphological and performance traits. Interestingly, high circulating androgens showed a nonlinear relationship with bite force in males and not females, with this relationship possibly being related to alternative male reproductive tactics. Our results suggest that androgens are not strongly correlated with most performance and morphological traits, although they may play an important organizational role during the development of morphological traits, which could explain the differences in morphology and thus performance between the sexes. Differences in performance between the sexes suggest differential selection on these functional traits between males and females. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 834–849.     

Reproductive skew and female trait elaboration in a cooperatively breeding rail

Intrasexual competition for reproduction is thought to be an important factor in the evolution of ornaments and weapons in males. However, the evolution of morphologically similar traits in females is often explained through other mechanisms, and the role of intrasexual competition in female trait elaboration has received little attention. Here, we explore the factors associated with female trait elaboration in the cooperatively breeding Pukeko (the New Zealand race of the Purple Swamphen Porphyrio porphyrio melanotus) by comparing sexual dimorphism in an ornament across two populations. Importantly, the two populations considered differ in several social factors that could affect the degree of female–female competition, and could thereby produce differential selection on elaborate female traits. Recent studies have suggested that high reproductive skew (i.e. monopolization of reproduction by dominant individuals) could influence the intensity of intrasexual competition and select for female elaboration. However, we found that sexual dimorphism was diminished and Pukeko females had more elaborate ornaments under conditions of low reproductive skew. We discuss alternative factors that could influence the degree of female–female competition, and show that reproductive skew may not always provide an accurate estimate of the scope for intrasexual competition.     

Exploring sexual differences in external morphology and limb muscles of Hylarana guentheri (Anura: Ranidae) during non-breeding season

Studying sexual dimorphism facilitates better understanding of the general intersexual divergence of the same species and gives insights into the impact of selective forces on each sex. The sexual dimorphism in anuran external morphology or limb muscles has been well studied in reproductive context, but less so outside the breeding season. Herein, the intersexual differences in 12 external morphological characteristics, 11 forelimb and 12 hindlimb muscles of Hylarana guentheri during non-breeding season were investigated. We found that the males possessed significantly greater head width and tympanum diameter, while the females had significantly larger body size and head length; there was no sexual differences in the examined limb muscles, except for the mass of flexor carpi radialis. The larger body size means greater fertility for the females, and the longer head may be related to the allometry and reduce resource competition. For the males, the wider head is likely correlated with prey size and male–male competition, and also, the larger tympanum and heavier flexor carpi radialis probably aid the productive success. This study provides the comprehensive morphological accounts about the sexual differences of H. guentheri during non-breeding season, which will contribute to clarify the sex-specific resource allocation and reproductive strategies of anurans.     

Sexual size dimorphism in a landlocked Pacific salmon in relation to breeding habitat features

Many animal species exhibit size dimorphism between sexes. Sexual selection, whereby male–male competition favors larger body sizes, has been considered a likely cause of sexual size dimorphism. Habitat features in breeding areas could affect the outcome of male–male competition, yet few attempts have been made to relate breeding habitat features with interpopulation variation in sexual size dimorphism. In this study, we examined interpopulation variation in sexual size dimorphism by studying the landlocked amago salmon (Oncorhynchus masou ishikawae) at a microgeographic scale. We found that female body size was independent of stream size but that male body size decreased with smaller stream sizes. A likely explanation is that the relationship between reproductive success and the size of males is influenced by the availability of refuges that are only available to small-bodied males. Sexual differences in body size increased with decreasing stream sizes, supporting the hypothesis that the reproductive success of larger males is reduced in smaller streams. In contrast, the maturation-length threshold increased with stream size for both sexes. The stream-size-based interpopulation variation in sexual size dimorphism and size at maturity in landlocked amago salmon may therefore have arisen through a combination of sexual and natural selection.     

Preliminary analysis of reproductive success in a large mammal with alternative mating tactics,the Northern chamois,Rupicapra rupicapra

In polygynous mating systems, reproductive skew depends on the ability of males to monopolize females, which in turn may promote the development of contrasting traits in the two sexes. Although dominant individuals normally enjoy a higher reproductive success (RS) than subordinates, the use of genetic markers has shown that behavioural observations of male mating success may not provide reliable clues of RS. We report the preliminary results of the first DNA‐based paternity analysis on the Northern chamois (Rupicapra rupicapra), a scarcely dimorphic mountain ungulate described as highly polygynous, in relation to mating tactic and age. Because of sampling difficulties, the success in parentage assignment was low, and the interpretation of results requires caution. Territorial males had a greater RS than nonterritorial ones but they were unable to monopolize mating events. Age had a weak effect on paternity outcome but only males ≥ 6 years showed siring success. Although future studies are needed to assess the opportunity for sexual selection in male chamois, the concurrence of limited sexual size dimorphism, compensatory growth, unbiased sex‐specific survival, RS of alternative mating tactics and, possibly, long breeding tenure, may hint at the adoption of a conservative mating strategy in this species.