Male-male interference competition decreases spawning rate in the European bitterling (<Emphasis Type="Italic">Rhodeus sericeus</Emphasis>)

We investigated the consequences of male-male interference competition associated with alternative male mating tactics in a freshwater fish, the European bitterling (Rhodeus sericeus). Male bitterling defend territories around living mussels and attract females to lay their eggs in the gill cavities of mussels. We experimentally manipulated spawning-site abundance and male density at two spatial scales. We showed that the total number of eggs spawned by females was constrained by the number of mussels available for oviposition. The effect was mediated by behavioral interactions among competing males because of variation in the Operational Sex Ratio (OSR) in close proximity to a mussel and not by a direct limitation in mussel capacity to accommodate the eggs. Both total and local male densities affected spawning behavior, and interacted in their effect on female spawning rate. Territorial male aggression caused courtship interruptions that prolonged the time until successful spawning and increased with male density. However, territoriality broke down at the highest male density, with a consequent stabilizing effect on spawning rate.Communicated by J. Krause     

Testes size and breeding systems in Japanese annrans with special reference to large testes in the treefrog,Rhacophorus arboreus (Amphibia : Rhacophoridae)

Summary Testes size was compared among 19 species of Japanese anurans in relation to their breeding systems. Although the mean body mass of the species examined varied markedly between 1.8 and 116 g, the mean proportion of testes mass to body mass was fairly constant at 0.2 to 0.4% across all species except the rhacophorid species. Foam-nest building rhacophorids had relatively large testes constituting more than 1% of their body mass. Among them, Rhacophorus arboreus had the largest, exceeding 5% of the body mass. Multi-male breeding, where a female is grasped by two or more males during spawning, occurs frequently in these rhacophorids, especially in R. arboreus. This close association between large testes size and multi-male breeding strongly suggests that sperm competition is an important factor affecting the evolution of relative testes size in Japanese anurans. Offprint requests to: T. Kusano     

Sperm competition and sexual selection: a meta-analysis of paternity studies of birds

Sperm competition (the competition among the sperm of different males for fertilization of the eggs of a female) has been suggested to be an important component of sexual selection, but no general assessment has been made of this proposition. We used a meta-analytic approach to assess the extensive literature on paternity (the proportion of offspring in a focal nest sired by an attending male) in birds based on allozyme and molecular techniques. The relative variance in male mating success was on average increased by a factor of 4.6 over the variance in apparent male success. Males with more extravagant secondary sexual characters had higher paternity in their own nests than less adorned males. There was a weak effect of male age being positively associated with paternity in own nests. Male body size measured as the length of wing and tarsus was weakly positively associated with paternity in own nests. Male survival prospect was positively associated with paternity in own nests. Polygynous males generally had decreased paternity of their broods compared to monogamous males. Paternity of the resident male decreased with increasing population density and breeding asynchrony. The intensity of paternity guards such as within-pair copulation rate and mate guarding were not significantly related to extra-pair paternity. Sperm competition was thus an important component of sexual selection by increasing the variance in male mating success, and by being associated with the expression of secondary sexual characters, in particular in dense and asynchronously breeding populations of birds. Received: 12 February 1998 / Accepted after revision: 31 May 1998     

Male-male competition ensures honest signaling of male parental ability in the three-spined stickleback (Gasterosteus aculeatus)

The importance of socially imposed costs for the evolution and maintenance of honest sexual signals has received less attention than other costs. Here I show that male-male competition can increase the honesty of sexual signaling in relation to male parental ability in a species with flexible signaling. When four three-spined stickleback (Gasterosteus aculeatus) males were allowed to court a female first separately and then in competition, red nuptial coloration under competition reflected male parental ability more accurately than red coloration when separated. Parental ability was determined as the ability of the male to raise a clutch of eggs to the hatching stage under interaction with other breeding and non-breeding males. The increased honesty under competition was probably due to subordinate males of poor parental ability decreasing their color expression under interaction to reduce the risk of fights with superior males. However, socially imposed costs of signaling were probably not the main factors maintaining honest signaling, as red coloration reflected male parental ability also in the absence of competition, although less accurately. Nevertheless, the small-scale differences that male-male competition induced can significantly facilitate adaptive female choice and have large impacts on sexual selection. Received: 7 July 2000 / Revised: 31 August 2000 / Accepted: 4 September 2000     

Male-male association patterns and female proximity in the guppy,Poecilia reticulata

If males differ in their ability to attract potential mates, and are able to perceive such differences, theory predicts they should distribute themselves in a manner that increases their probability of obtaining potential matings. The relationship between male-male association patterns and the proximity of females in social groups, however, remains virtually unexplored. Experimental analysis of this relationship in the guppy, Poecilia reticulata, demonstrates that in preference tests males showed a strong tendency to associate with other males that were further away from potential mates than they were themselves. Male guppies pursue a behavioral strategy that involves categorizing other males based on their proximity (and possibly relative attractiveness) to females, remembering the identity of such individuals, and using this information when choosing between other males as associates. Such a strategy may increase a male's chances of being the individual chosen by a female assessing nearby males.     

Body size and sexual selection in the koala

Sexual selection is often characterized by polygynous breeding systems, size dimorphism, and skewed operational sex ratios. Koalas are sexually dimorphic in multiple domains, yet are absent from the literature on sexual selection and the structure of their mating system is unclear. We provide the first documentation of the strength of sexual selection in koalas by using microsatellite markers to identify sires. We combine the genetic data with morphological data in order to assess the role of body size in regulating reproductive output. During our 4-year study, 37% of males were identified as possible sires. Males were significantly larger than females, with sires heavier than non-sires. Male body mass correlated with annual reproductive output, with Crow’s Index of Opportunity for Selection revealing that variation in male reproductive success was threefold higher than that of females. Since it appears that male koalas rarely engage in physical confrontations over access to females, size dimorphism could be based upon non-agonistic competition and/or female mate choice. We propose that size dimorphism in koalas evolved as a consequence of endurance rivalry promoting vocal sexual advertisements that attract females. We suggest that female choice is a key mediator of male reproductive output.     

Phylogenetic analyses of sexual selection and sexual size dimorphism in pinnipeds

Pinnipedia contains some of the most spectacular examples of sexual size dimorphism, examples that are therefore frequently used to illustrate the theory of sexual selection. This paper addresses the question of whether a significant relationship between sexual selection and size dimorphism exists in a comparative context. Thus, harem size and body size data gathered from the literature were analysed with independent contrasts analyses. These investigations showed that sexual size dimorphism is not a consequence of an allometric relationship between male and female size. Instead, there is a clear relationship between harem size and sexual size dimorphism. Further analyses also revealed a significant relationship between harem size and male size whereas no such relationship existed for females. These results support the hypothesis that sexual size dimorphism in pinnipeds is the product of an exclusively male response to sexual selection.     

Sexual conflict,sexual selection and sperm competition in the spawning decisions of bitterling, <Emphasis Type="Italic">Rhodeus</Emphasis><Emphasis Type="Italic">sericeus</Emphasis>

Sexual selection may operate through either direct selection on preference genes or indirect genetic benefits. However, in many species both direct and indirect selection may operate and can influence female mate and oviposition choice. Adaptations by males for sperm competition can also affect female mate and oviposition choice and can lead to sexual conflict. We investigated the role of direct and indirect benefits and male adaptation to sperm competition for female oviposition decisions in the bitterling, Rhodeus sericeus, a freshwater fish that spawns on the gills of living unionid mussels. Field experiments showed that females chose mussels for spawning on the basis of the number of embryos already in mussels and the body size and/or extent of red coloration of the eyes of territorial males. There was also a significant interaction between these direct and indirect benefits of oviposition choice. Territorial male bitterling avoided leading females to mussels into which rivals had already released sperm. Males also increased their rate of sperm release into mussels in response to sperm release by sneaking males. We discuss the possibility that sexual conflict in oviposition choice occurs in this species.     

Advertisement-call modification,male competition,and female preference in the bird-voiced treefrog <Emphasis Type="Italic">Hyla avivoca</Emphasis>

Senders and receivers influence dynamic characteristics of the signals used for mate attraction over different time scales. On a moment-to-moment basis, interactions among senders competing for a mate influence dynamic characteristics, whereas the preferences of receivers of the opposite gender exert an influence over evolutionary time. We observed and recorded the calling patterns of the bird-voiced treefrog Hyla avivoca to assess how the dynamic characters of calls vary during interactions among groups of males in a chorus. This question was also addressed using playback experiments with males. Playback experiments with females showed how changes in dynamic call properties are likely to affect male mating success. Frogs calling in pairs, groups, or in response to playbacks produced longer calls than did isolated males. During call overlap, males often increased the duration of the silent interval (gaps) between the pulses of their calls so that the pulses of the calls of two neighbors interdigitated. This change resulted in increased variability of pulse rate, a traditionally static acoustic property; however, males also produced high proportions of non-overlapped calls in which variability in pulse rate was low and had species-typical values. Females preferred long calls to short- and average-duration calls, and non-overlapped calls to overlapped calls. Given a choice between pairs of overlapped calls, females preferred pairs in which the proportion of overlap was low and pairs in which the pulses of such calls interdigitated completely. The observed patterns of vocal competition thus reflect the preferences of conspecific females, which have influenced the evolution of the calling behavior of H. avivoca. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Pre-copulatory sexual selection in the cigarette beetle Lasioderma serricorne

Traditional concepts of sexual selection and sexual conflict make different predictions about the costs and benefits to females of exposure to males with higher mating success. The traditional concepts of sexual selection assume that females benefit from their mate choices, whereas sexual conflict assumes that the females suffer greater costs by mating with males who have greater mating success and thus reduce their fitness. In order to understand how mate choice evolves, it is necessary to estimate the overall effect of mate choice on female fitness. However, relatively few studies have conducted that investigation. In this study, we investigated the direct and indirect effects of mating with attractive males on the fitness of females in the cigarette beetle Lasioderma serricorne. Mating with attractive males increased the number of female offspring but did not affect female longevity. Additionally, we found evidence that attractive males sire highly attractive sons. Thus, mating with an attractive male provides direct and indirect benefits but no fitness cost to female L. serricorne.     

Bower destruction and sexual competition in the satin bowerbird (Ptilonorhynchus violaceus)

Summary Male satin bowerbirds often destroy the bowers of other males. Bowers are a key element in male sexual display and their destruction represents a unique pattern of sexual competition. For two mating seasons bowers of displaying males were continously monitored to produce a complete record of bower destructions. The number of destructions at bowers and the amount of destruction of bowers were invesely correlated with bower quality. The best predictor of male bower destroying behaviour is male aggressiveness at feeding sites. Males directed most destructions at adjacent bower sites. These results show that male interactions are important in determining differences in the quality of display among male satin bower-birds, and are consistent with the view that females use bowers as indicators of male quality in mate choice.     

Male asymmetry and postcopulatory sexual selection in the fly Dryomyza anilis

The significance of male asymmetry in postcopulatory sexual selection was studied in the fly Dryomyza anilis by examining whether male asymmetry is related to fertilization success. The traits measured were wing length, tibia length and the length of small and large claspers. The male claspers are situated at the tip of the abdomen, functional pair of claspers consists of a large and a small clasper on the same side of the body. These claspers are used to tap the female abdomen after sperm transfer, which has been shown to increase fertilization success for the mating male. Fertilization success was negatively related to the fluctuating asymmetry of wing length, suggesting either female preference for more symmetrical males or a relationship between male asymmetry and intrasexual selection which was reflected in mating performance. Fertilization success was also related to the length of small claspers, decreasing with increasing length of the claspers. In addition, males with asymmetrical small claspers enjoyed higher fertilization success than symmetrical ones. This study shows that fluctuating asymmetry in wing length is an important fitness trait in postcopulatory sexual selection. Since male tapping affects sperm distribution in the female's sperm storage organs, the higher fertilization success of males with asymmetrical small claspers could have a functional relationship with the asymmetrical position of female sperm storage organs. Received: 6 March 1997 / Accepted after revision: 8 November 1997     

Predation, scramble competition, and the vigilance group size effect in dark-eyed juncos (Junco hyemalis)

In socially feeding birds and mammals, as group size increases, individuals devote less time to scanning their environment and more time to feeding. This vigilance “group size effect” has long been attributed to the anti-predatory benefits of group living, but many investigators have suggested that this effect may be driven by scramble competition for limited food. We addressed this issue of causation by focusing on the way in which the scan durations of free-living dark-eyed juncos (Junco hyemalis) decrease with group size. We were particularly interested in vigilance scanning concomitant with the handling of food items, since a decrease in food handling times (i.e. scan durations) with increasing group size could theoretically be driven by scramble competition for limited food resources. However, we showed that food-handling scan durations decrease with group size in an environment with an effectively unlimited food supply. Furthermore, this food-handling effect was qualitatively similar to that observed in the duration of standard vigilance scans (scanning exclusive of food ingestion), and both responded to changes in the risk of predation (proximity of a refuge) as one might expect based upon anti-predator considerations. The group size effects in both food-handling and standard scan durations may reflect a lesser need for personal information about risk as group size increases. Scramble competition may influence vigilance in some circumstances, but demonstrating an effect of competition beyond that of predation may prove challenging. Received: 22 September 1998 / Received in revised form: 1 February 1999 / Accepted: 14 February 1999     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

Female egg investment in relation to male sexual traits and the potential for transgenerational effects in sexual selection

Life-history theory predicts that individuals should increase their reproductive effort when the fitness return from reproduction is high. Females mated with high-quality males are therefore expected to have higher investment than females mated with low-quality males, which could bias estimates of paternal effects. Investigating the traits females use in their allocation decisions and the aspects of reproduction that are altered is essential for understanding how sexual selection is affected. We studied the potential for differential female allocation in a captive population of a precocial bird, the Chinese quail, Coturnix chinensis. Females paired with males with large sexual ornaments laid larger, but not more, eggs than females paired with males with small sexual ornaments. Furthermore, female egg mass was also significantly positively affected by male testis size, probably via some unknown effect of testis size on male phenotype. Testis size and ornament size were not correlated. Thus, both primary and secondary male sexual traits could be important components of female allocation decisions. Experimental manipulation of hormone levels during embryonic development showed that both male and female traits influencing female egg size were sensitive to early hormone exposure. Differences in prenatal hormone exposure as a result of maternal steroid allocation to eggs may explain some of the variation in reproductive success among individuals, with important implications for non-genetic transgenerational effects in sexual selection.Communicated by C. Brown     

Sexual selection and cuckoldry in a monogamous songbird: implications for sexual selection theory

Sexual selection is generally assumed to be weaker in monogamous than in polygynous animals. Recently, though, extra-pair fertilizations have been hailed as an important force in generating variance in reproductive success among males in socially monogamous species, thereby increasing the intensity of sexual selection. To see if extra-pair copulations contribute to variance in male reproductive success in the house finch (Carpodacus mexicanus), we used DNA fingerprinting to determine the paternity of chicks from 35 nests. This species is a socially monogamous passerine in which plumage brightness serves as a sexually selected indicator of male quality. Out of 119, nestlings 10 (8.3%) were fathered by a male other than the attending male, but cuckoldry occurred randomly with respect to the plumage colouration, size, or age of the attending male. Thus extra-pair fertilizations do not generate variance in male reproductive success with respect to plumage colour. On the other hand, a strongly male-biased sex ratio and asynchronous breeding by females may generate substantial variance in male reproductive success and could explain the evolution of ornamental colouration.     

Male-male competition selects for delayed sex change in the protandrous shrimp<Emphasis Type="Italic"> Pandalus latirostris</Emphasis>

In most protandrous species, male size advantage is generally regarded as unimportant in determining the timing of protandrous sex change. In pandalid shrimp, the size/age of male sex change often fluctuates among years and populations, but the adaptive significance of late reversing males (LRMs) is not well understood. This study experimentally examined the adaptive significance of LRMs in the protandrous pandalid shrimp Pandalus latirostris Rathbun. Field and laboratory studies were carried out in August–September of 1998–2002 on P. latirostris in Notoro Lagoon, Japan (44°03′N; 144°10′E). Mature females that had molted (i.e. mate receptive) were tethered in the field and their mating behavior with wild males was observed. Copulations occurred with a single male at a time, although other males could sequentially mate with a tethered female. Because tethered females rejected male approaches, males had difficulty transferring their spermatophores. In the laboratory, males copulated with non-tethered, recently molted females for only 15 min after molting. Recently molted females are wary of potential predators, since their soft exoskeleton makes them particularly vulnerable. Fast access by males enhanced fertilization success in this shrimp. The effect of male size on mating success in the laboratory was examined. Both small and large males successfully inseminated females in the absence of competitors. In experiments where large, medium, and small males competed for a female, however, larger males guarded females longer than smaller males, until the female molted and became receptive. Moreover, large males were more successful at copulating once molting occurred. These results imply that male-male competition drives delayed sex change in some situations.     

Multiple spawning events and sexual reproduction in the octocoral Sarcophyton elegans (Cnidaria: Alcyonacea) on Lizard Island, Great Barrier Reef

Sarcophyton elegans is a common symbiotic (zooxanthellate) octocoral species in the shallow waters of the Great Barrier Reef (GBR). Study of a population at Lizard Island (14°40′S, 145°28′E) on the GBR from October 1991 to January 1994 revealed that, as is typical of tropical alcyonarian corals, S. elegans is a gonochoric broadcast spawner with a 1:1 sex ratio. Sexual reproduction was closely correlated with colony size, with first reproduction at 13-cm basal stalk circumference for females and 12 cm for males. Oogenesis took 19–24 months, with a new cycle commencing every year, and spermatogenesis took 10–12 months. The majority of gametes were released during the annual austral mass coral spawning event after the full moon in November, but gametes were also released after the full moon in each month between August and February. All autozooid polyps participated in reproduction, but those at the outer edge of a colony released their gametes first. During subsequent months, the polyps closer to the center of the colony released their gametes. This is a novel strategy of gamete release, reported here for the first time, which accommodates the demands of feeding and reproduction in a different way than other corals where individual polyps have separate feeding or reproductive roles. Colonies upstream in the prevailing current spawned up to 1 month earlier than those downstream and ceased 1 month earlier. The mechanism controlling this spatial differentiation in spawning time, repeatedly observed over three seasons, is unknown. Sarcophyton elegans appears to have a dual strategy of providing protection for its gametes by releasing most of them concurrently with the single, annual mass spawning of a large number of cnidarians, while also hedging its bets by individual colonies spawning a fraction of their gametes over an extended period of 6 months.