Male–male competition for large nests as a determinant of male mating success in a Japanese stream goby, Rhinogobius sp. DA

Males of the stream goby Rhinogobius sp. DA (dark type) court females in deep pools and care for the eggs under stones in shallow riffles. We studied male–male competition for access to females and nest sites to understand how male size influences the mating success of this species. In field observations, larger males won in fighting with other males. However, large males did not tend to monopolize courtship opportunities, and the frequency of successful courtships, after which males led the females to the nests, was not related to male body size. The fact that courted females always escaped from the fighting sites once males began fighting likely explains why male size was not positively related to courtship success. Large males occupied large nest stones, and the number of eggs received in the nest was correlated positively with nest size. In aquarium experiments with two tiles of different sizes provided as nesting materials, males always chose the larger nest and, when two males were introduced simultaneously, the larger one occupied the larger nest. These results suggested that male mating success of this goby is determined by male–male competition for large nests rather than for access to females. Received: June 9, 2000 / Revised: September 2, 2000 / Accepted: October 4, 2000     

Females decide whether size matters: plastic mate preferences tuned to the intensity of male-male competition

Recent studies have indicated that mating success of large malesmay improve under increasing levels of mating competition. Thisoutcome is explained 1) if male mating competition is overridingfemale preferences for male traits that are unrelated to, ornegatively correlated with, male size and dominance and, inso doing, dictates the distribution of matings or 2) if femalesalter their preferences with respect to large males when male–malecompetition is intense. Under both hypotheses, one could expectlarge, dominant males to be more successful under intense competitionthan under weak competition. However, only the first explanationpredicts that male mating success under intense competitionshould be determined by dominance; traits that are unrelatedto male dominance should be uncorrelated to mating success.In contrast, if females change their preferences (explanation2), males with traits beneficial to females independent of thecompetitive environment can maintain a high mating success underall levels of male–male competition. We tested these alternativesusing a small marine fish, the sand goby, Pomatoschistus minutus.The mating success of large males increased under conditionsallowing intense male competition, whereas females showed apreference for good nest building independent of the level ofcompetition. These findings suggest that females are in controlof their choice by altering their preference for male size inresponse to the intensity of male–male competition ratherthan female preference being overshadowed by male dominance.This plasticity of preferences implies that the strength ofsexual selection is not constant at the population level.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

Relatively large males lower reproductive success in female zebrafish

Females can adjust their reproductive effort in relation to their partner’s perceived fitness value. In zebrafish (Danio rerio), large males are typically preferred mating partners. However, females have been observed to reduce their reproductive output with exceptionally large males but it remains unknown whether it is due to sexual harassment or aggressive behavior to establish and maintain dominance. Here, we study the association between relative male size, sexual harassment and dominance behavior, female stress status (stress behaviors and whole-body cortisol concentration), and reproductive success during a 4 day spawning trial. We found female cortisol to correlate negatively with female body size and positively with female dominance behavior. However, male and female behavior as well as female cortisol level were not related to relative male size. Females mating with relatively large males produced more and most of their eggs during the first spawning day, while females with smaller males produced few eggs during the first day but then increased egg production. Despite females produced more eggs when mating with relatively larger males, their eggs had substantially lower fertilization rates compared to females mating with relatively smaller males. Hence, overall, the reproductive fitness was lowest when females mated with a relatively large male. These findings could help to explain the maintenance of male size variation under natural conditions.     

Determinants of Paternity Success in a Group of Captive Vervet Monkeys (<Emphasis Type="BoldItalic">Chlorocebus aethiops sabaeus</Emphasis>)

Paternity success of high-ranking primate males is affected by the number of males and the number of fertile females and their cycle synchrony. Female vervets in the wild show strong reproductive seasonality and do not advertise the ovulatory period with conspicuous signals or behavior. Because this makes it difficult for males to monopolize fertile females, it can be expected that male reproductive skew in this species is lower than in other cercopithecines living in multimale groups that advertise the ovulatory period. We assessed male reproductive success in a captive vervet group, initially consisting of 4 males and 12 unrelated females. Besides a general low reproductive skew, we predicted paternity success of the alpha males to be dependent on the overlap of synchronously fertile females, month into alpha male tenure, and housing conditions (the subjects were kept in a large park but had to be locked in a small indoor compartment during the winter months). Further, because the number of males reaching their prime increased over time, we predicted a decrease in reproductive success of the alpha male with increasing tenure length of the alpha male. To assess this we collected genetic samples from 27 of 32 offspring born into the group during 3 yr, using 7 polymorphic human microsatellite markers. Contrary to our predictions, reproductive success of the males occupying the alpha position during the study was among the highest found in cercopithecines, with the alpha males siring 78% of all offspring. The degree of overlapping fertile periods did not affect paternity success, nor did paternity success of the alpha male differ between the 2 housing conditions. Only the alpha male’s tenure length had an effect. Alpha male reproductive success decreased over time, which we attribute to either a shift in female mate choice toward younger males or to increased competition among the males in the later stage of the alpha male’s tenure. This study demonstrates the importance of assessing reproductive success of males throughout >1 season and should spawn more research into the role of female choice and postcopulatory mechanisms in this species.     

Emergence and mating behavior of the oriental fruit moth Cydia molesta (Lepidoptera: Tortricidae) and its potential for reproduction

The Oriental fruit moth, Cydia molesta (Busck, 1916) (Lepidoptera: Tortricidae), is a key pest of fruit and is widely distributed around the world. There are important connections between its behavior and biology and its management in agriculture, but few studies have investigated the associations between adult behaviors and oviposition. In this study, adult emergence, mating, and reproduction were investigated under laboratory and field conditions. The ratio of females to males at eclosion was approximately 1:1. When one virgin female had access to one virgin male, 66% and 34% of the couples copulated just once and twice, respectively; and the infertility rate of eggs (21.39 ± 1.25%) did not vary daily. Males, given access to one new female daily, could copulate multiple times, whereas females seldom mated more than once, indicating a male-biased operational sex ratio, but mating status of the male parent had no effect on progeny egg reproduction. Also, the number of eggs that hatched by all female partners of a male was inversely proportional to copulation duration for the female laying the eggs for total female reproductive success; and the number of eggs laid by all female partners of a male was proportional to their number of matings for total male reproductive success. However, the total number of eggs that hatched did not significantly differ for eggs laid by a female given new virgin males daily for mating (17.75 ± 4.28) versus eggs laid by virgin females (19.17 ± 7.51) presented daily with a male that re-mated daily with the series of females. Therefore, our results showed that females engaged in mate choice and males engaged in mate competition, affecting egg production, a factor that may be used to enhance mating disruption technology against Cydia molesta.     

Fitness consequences of male provisioning of incubating females in a desert passerine bird

Male provisioning of incubating females can increase reproductive success by maintaining physiological condition of females and consistency of incubation. The effects of male provisioning on the maintenance of incubation temperature and embryo development should be particularly pronounced in environments where ambient temperature exceeds the tolerance of unincubated eggs and where consistency of female incubation might be particularly important for hatching success. Here, we investigated the reproductive consequences of incubation feeding in a desert population of House Finches (Carpodacus mexicanus) in southwestern Arizona. We found that greater nest attentiveness by females was related to higher minimum incubation nest temperature, that in turn was closely associated with hatching success. Only 44% of males regularly provisioned their incubating females. Although provisioned females maintained higher incubation temperature and took fewer incubation breaks than non-provisioned females, overall, male provisioning did not influence incubation dynamics or hatching success. Further, a male’s incubation feeding rate did not correlate with male provisioning of nestlings. These results corroborate the finding that, in male House Finches, neither provisioning of incubating females nor pre-incubation courtship feeding are associated with increases in circulating pituitary prolactin––the hormone regulating male provisioning of nestlings. We suggest that incubation provisioning by male might be a component of pair maintenance behavior and that variation in male incubation behavior is best understood in relation to asymmetries in residual reproductive values between the mates.     

Female choice and male reproductive success in the angel blenny,Coralliozetus angelica (Teleostei: Chaenopsidae)

Correlates of reproductive success of male Coralliozetus angelica were studied in field populations in the central Gulf of California by collecting males with their shelters (vacant barnacle tests) and counting the number of eggs defended. Male body size was positively correlated with the number of eggs defended. This apparently resulted from female preference for large males since females preferred to mate with the larger of two males in laboratory choice experiments. Condition of males, as assessed by their relative head width, was also positively correlated with the number of eggs defended. Fit of males in their shelter entrance, male colour intensity, and male size relative to immediately surrounding males were not correlated with reproductive success. The quality of shelters was an important determinant of male reproductive success since females apparently avoided mating with males in heavily fouled shelters. Large shelters tended to be more heavily fouled than small shelters and male size and shelter size were positively correlated. The resulting negative correlation of male size and shelter quality apparently constrained female preference for large males and may be an important factor limiting the degree of polygyny in this species.     

Reproductive regulation in the female Japanese minnow, Pseudorasbora parva (Cyprinidae)

The Japanese minnow, Pseudorasbora parva, is a multiple spawner that lays eggs repeatedly during the spawning season. In laboratory aquaria, 19 of 63 females did not spawn, whereas 44 other females laid 167 to 6285 eggs in 1 to 14 mating sequences. Neither fish density nor sex ratio (number of males per female) affected fecundity, but the increase in fish density significantly decreased the growth rate of females. The increase in male length significantly increased fecundity in aquaria which contained only a single male, whereas it decreased female growth rate. These relationships were not evident in aquaria housing three or ten males. Since larger males were more dominant and had larger testes than smaller males, the correlation between fecundity and male size strongly suggests that individual females regulated fecundity to increase their reproductive success.     

Indirect female choice mediated by sex pheromones in the hermit crab Pagurus filholi

Males of the hermit crab Pagurus filholi perform precopulatory guarding behavior, and solitary males often show aggressive behavior to take away guarded females. Males behave coercively while guarding females, so direct mate choice by females seems difficult in such a situation. By performing several experiments we examined possible indirect female choice of hermit crab. Males were attached to a shell by their left cheliped to look like guarding pairs (fake guarding pairs). The shells were filled with cotton containing either seawater or pheromone water. The fake guarding pair with only seawater caused male–male combat in 60% of trials whereas with pheromone water combats occurred in 88% of trials. Mean duration of male–male combat was significantly longer in trials with drops of seawater containing pheromones than in those without pheromones. These results suggest guarding pairs themselves cause male–male combat by visual stimulation, that female sex pheromones have further significant function in the recognition of guarding pairs and intensification of male–male combat, and that females release sex pheromones while they are guarded. As a result of the combat, the larger male ended up guarding a female. This strongly suggests that females choose males indirectly by exploiting male–male competition induced by sex pheromones under male coercive behavior.     

Overt and covert competition in a promiscuous mammal: the importance of weaponry and testes size to male reproductive success

Male contests for access to receptive females are thought to have selected for the larger male body size and conspicuous weaponry frequently observed in mammalian species. However, when females copulate with multiple males within an oestrus, male reproductive success is a function of both pre- and postcopulatory strategies. The relative importance of these overt and covert forms of sexual competition has rarely been assessed in wild populations. The Soay sheep mating system is characterized by male contests for mating opportunities and high female promiscuity. We find that greater horn length, body size and good condition each independently influence a male's ability to monopolize receptive females. For males with large horns at least, this behavioural success translates into greater siring success. Consistent with sperm-competition theory, we also find that larger testes are independently associated with both higher copulation rates and increased siring success. This advantage of larger testes emerges, and strengthens, as the number of oestrous females increases, as dominant males can no longer control access to them all. Our results thus provide direct quantitative evidence that male reproductive success in wild populations of mammals is dependent upon the relative magnitude of both overt contest competition and covert sperm competition.     

Mating frequency and reproductive success in an income breeding moth,Mnesampela privata

The frequency of mating in insects is often an important determinant of female reproductive output and male sperm competition. In Lepidoptera that provide male nutrients to the female when mating, it is hypothesized that polyandry may be more prevalent. This is thought to be especially so among species described as income breeders; that is, in species who do not derive all their nutrients for reproductive output entirely from the resources obtained during the larval stage. We selected the geometrid moth, Mnesampela privata (Guenée) (Lepidoptera: Geometridae), to examine this hypothesis further. We found this species was best characterized as an income breeder with female weight on emergence positively correlated with total egg load but not with the number of eggs laid. Further, in accord with income breeders, females emerged with a partially developed egg load and lifetime fecundity was positively correlated with the number of oviposition days. However, in the laboratory we found that incidence of repeated matings or polyandry was rare. When moths were paired singly over their lifetime, only 4% of mated females multiple mated. When females were paired with three males concurrently, female mating success increased from 60 to 81% with multiple mating among mated females increasing to just 15%. Dissection of wild caught M. privata found that polyandry levels were also low with a maximum of 16.4% of females collected at any one time being multiple mated. In accord with theory, mating significantly increased the longevity of females, but not of males, suggesting that females acquire essential resources from male ejaculates. Despite this, multiple mated females showed a trend toward decreasing rather than increasing female reproductive output. Spermatophore size, measured on death of the female, was not correlated with male or female forewing length but was negatively correlated with the number of fertile eggs laid and female longevity. Smaller spermatophore width may be related to uptake of more nutrients by the female from a spermatophore. We discuss our findings in relation to income breeding and its relationship to polyandry in Lepidoptera.     

Size-assortative pairing in the lotic amphipod <Emphasis Type="Italic">Gammarus zaddachi</Emphasis>, an examination of hypotheses and the influence of current speed

Three main hypotheses have been put forward to explain size-assortative pairing in gammarid amphipods: microhabitat separation, sexual selection and loading constraint. In order to determine which hypothesis best explains this phenomenon in the estuarine species Gammarus zaddachi, I first measured the body lengths and dry weights of precopula pairs collected from two field sites with substantially different current speeds. Second, I performed three laboratory experiments in order to estimate the importance of the following processes: (1) male choice; (2) male–male competition and (3) male–female acceptability. The loading constraint hypothesis seemed best supported by the data in that field-collected male G. zaddachi size correlated well with female size in precopula pairing in both fast and slow flowing water. In the laboratory, males preferred females of their same size group (large versus small), and ‘won’ them in the male–male competition experiments. Size-assortative pairing is thus likely a consequence of the loading constraints imposed upon these males by virtue of them having to carry and manoeuvre their partners through flowing water, while attempting to maintain station in an optimal microhabitat. Males may therefore forego the largest, most fecund females, in favour of a practicable payload (small male–large female pairings were rare). However, there seems to be a lower limit to this selection, indicated by the high degree of cannibalism on small females by large males.     

Reproductive ecology of the jacky dragon (Amphibolurus muricatus): an agamid lizard with temperature‐dependent sex determination

Abstract The jacky dragon, Amphibolurus muricatus (White, ex Shaw 1790) is a medium sized agamid lizard from the southeast of Australia. Laboratory incubation trials show that this species possesses temperature‐dependent sex determination. Both high and low incubation temperatures produced all female offspring, while varying proportions of males hatched at intermediate temperatures. Females may lay several clutches containing from three to nine eggs during the spring and summer. We report the first field nest temperature recordings for a squamate reptile with temperature‐dependent sex determination. Hatchling sex is determined by nest temperatures that are due to the combination of daily and seasonal weather conditions, together with maternal nest site selection. Over the prolonged egg‐laying season, mean nest temperatures steadily increase. This suggests that hatchling sex is best predicted by the date of egg laying, and that sex ratios from field nests will vary over the course of the breeding season. Lizards hatching from eggs laid in the spring (October) experience a longer growing season and should reach a larger body size by the beginning of their first reproductive season, compared to lizards from eggs laid in late summer (February). Adult male A. muricatus attain a greater maximum body size and have relatively larger heads than females, possibly as a consequence of sexual selection due to male‐male competition for territories and mates. If reproductive success in males increases with larger body size, then early hatching males may obtain a greater fitness benefit as adults, compared to males that hatch in late summer. We hypothesize that early season nests should produce male‐biased sex ratios, and that this provides an adaptive explanation for temperature‐dependent sex determination in A. muricatus.     

The Effect of Female Quality on Male Ejaculatory Expenditure and Reproductive Success in a Praying Mantid

Strategic ejaculation is a behavioural strategy shown by many animals as a response to sperm competition and/or as a potential mechanism of cryptic male choice. Males invest more mating resources when the risk of sperm competition increases or they invest more in high quality females to maximize their reproductive output. We tested this hypothesis in the false garden mantid Pseudomantis albofimbriata, where females are capable of multiply mating and body condition is an indicator of potential reproductive fitness. We predicted male mantids would ejaculate strategically by allocating more sperm to high quality females. To determine if and how males alter their ejaculate in response to mate quality, we manipulated female food quantity so that females were either in good condition with many eggs (i.e. high quality) or poor condition with few eggs (i.e. low quality). Half of the females from each treatment were used in mating trials in which transferred sperm was counted before fertilisation occurred and the other half of females were used in mating trials where fertilisation occurred and ootheca mass and total eggs in the ootheca were recorded. Opposed to our predictions, the total number of sperm and the proportion of viable sperm transferred did not vary significantly between female treatments. Male reproductive success was entirely dependent on female quality/fecundity, rather than on the number of sperm transferred. These results suggest that female quality is not a major factor influencing postcopulatory male mating strategies in P. albofimbriata, and that sperm number has little effect on male reproductive success in a single mating scenario.     

Identifying a causal agent of sexual selection on weaponry in an insect

In many animal species, males do not seek females directly butinstead locate and defend sites that contain spatially or temporallylimited resources essential to female survival and reproduction.Resident males that successfully repel conspecific rivals canmate with females attracted to these resources. In theory, increasingresource value increases harem size and thus increases the opportunity(I_mates) for and strength of sexual selection on traits crucialto male resource-holding potential and mating success. I experimentallytested this hypothesis in the field using the Wellington treeweta, Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae),a sexually dimorphic insect in which males use their enlargedmandibles as weapons in male–male contests over accessto females sheltering in tree cavities (galleries). By manipulatinggallery size, I showed that, compared with smaller galleries,larger galleries housed larger harems. Variation in gallerysize was an important determinant of I_mates, but contrary toexpectation, greater opportunity existed in small galleriescompared with large galleries. As predicted, male weapon sizewas under stronger directional selection in large galleriesbecause the fitness benefits were greater under these conditionscompared with small galleries. My results help explain the positiveassociation between average weapon size and average gallerysize observed within and among tree weta populations in NewZealand.     

Group size and composition influence male and female reproductive success in black howler monkeys (Alouatta pigra)

It has been argued that grouping patterns might influence the reproductive performance of individuals. Increasing group size results in greater travel costs and competition over depletable food resources, which could lead to reduced individual reproductive success. However, in groups with an increasing number of males, female reproductive success is predicted to augment because larger male groups might better protect immatures from infanticidal attacks. In contrast, male reproductive success is predicted to decrease with number of males in a group because fertilization cannot be shared between males. In this paper, we test these predictions on the Mesoamerican black howler monkey (Alouatta pigra) with data on group size and composition for 120 groups from eight populations of black howler monkeys existing in eight protected forests in Mexico and Guatemala. Male and female reproductive success were calculated as a deviation of the observed number of infants (or immatures) from the expected number of infants (or immatures), relative to the number of males and females in a group. Results indicate that both male and female reproductive success decreased with group size. Male reproductive success decreased with an increasing number of males in a group and with increasing proportion of males relative to females in a group. Decreased female reproductive success was associated with increasing number of females in a group, and female reproductive success had a tendency to increase with increasing number of males in a group. These results suggest that in black howler monkeys, living in larger groups might negatively affect the reproductive success of each member. Our findings are similar to those reported for a population of a sister species, Alouatta palliata, living in larger groups.     

Social network predicts loss of fertilizations in nesting males of a fish with alternative reproductive tactics

Alternative reproductive tactics (ARTs) evolve when there is strong intra-sexual competition between conspecifics for access to mates. Typically, larger “bourgeois” males reproduce by securing the access to reproductive resources while smaller “parasitic” males reproduce by stealing fertilizations from larger males. A number of factors can influence the reproductive success of each tactic, including intrinsic (e.g. size) and extrinsic (e.g. tactic relative frequency) variables. An example where plastic ARTs occur is the peacock blenny Salaria pavo, with large males reproducing by defending nests and attracting females (bourgeois tactic) and small males reproducing by achieving sneaked fertilizations (parasitic tactic). In this study, we conducted field observations on individually tagged animals to determine their social network and collected eggs from 11 nests to determine the fertilization success of each male tactic. Paternity estimates for 550 offspring indicated an average fertilization success for nest-holder males of 95%. Nest-holder male morphological traits and social network parameters were tested as predictors of fertilization success, but only the number of sneakers present in the nest-holder’s social networks was found to be a predictor of paternity loss. Although male morphological traits had been previously found to be strongly correlated with reproductive success of nest-holder males, as measured by the number of eggs collected in the male’s nest, no correlation was found between any of the measured morphological traits and fertilization success for these males. The results suggest a stronger influence of the social environment than of morphological variables in the proportion of lost fertilizations by nest-holder males of this species.     

Paternity costs from polyandry compensated by increased fecundity in the hide beetle

Polyandry-induced sperm competition is assumed to impose costson males through reduced per capita paternity success. In contrast,studies focusing on the consequences of polyandry for femalesreport increased oviposition rates and fertility. For thesespecies, there is potential for the increased female fecundityassociated with polyandry to offset the costs to males of sharedpaternity. We tested this hypothesis by comparing the proportionand number of offspring sired by males mated with monandrousand polyandrous females in the hide beetle, Dermestes maculates,both for males mating with different females and for males rematingwith the same female. In 4 mating treatments, monandrous femalesmated either once or twice with the same male and polyandrousfemales mated either twice with 2 different males or thricewith 2 males (where 1 male mated twice). Polyandrous and twice-matingmonandrous females displayed greater fecundity and fertilitythan singly mating monandrous females. Moreover, males rematedto the same female had greater paternity regardless of whetherthat female mated with another male. In both polyandrous treatments,male mating order did not affect paternity success. Finally,although the proportion of eggs sired decreased if a male matedwith a polyandrous female, multiply mating females or femalesthat remated with a previous mate laid significantly more eggsand thus the actual number of eggs sired was comparable. Thus,males do not necessarily accrue a net fitness loss when matingwith polyandrous females. This may explain the absence of anyobvious defensive paternity-protection traits in hide beetlesand other species.     

Sexual conflict over mating in Gnatocerus cornutus? Females prefer lovers not fighters

Female mate choice and male–male competition are the typical mechanisms of sexual selection. However, these two mechanisms do not always favour the same males. Furthermore, it has recently become clear that female choice can sometimes benefit males that reduce female fitness. So whether male–male competition and female choice favour the same or different males, and whether or not females benefit from mate choice, remain open questions. In the horned beetle, Gnatocerus cornutus, males have enlarged mandibles used to fight rivals, and larger mandibles provide a mating advantage when there is direct male–male competition for mates. However, it is not clear whether females prefer these highly competitive males. Here, we show that female choice targets male courtship rather than mandible size, and these two characters are not phenotypically or genetically correlated. Mating with attractive, highly courting males provided indirect benefits to females but only via the heritability of male attractiveness. However, mating with attractive males avoids the indirect costs to daughters that are generated by mating with competitive males. Our results suggest that male–male competition may constrain female mate choice, possibly reducing female fitness and generating sexual conflict over mating.