No reason to sneak: why males of all sizes can breed in the hole-nesting blenny, Aidablennius sphinx

Most blennioid fishes show a resource-based, promiscuous mating system with alternative 'sneaking' mating tactics. The hole-nesting species, Aidablennius sphinx, however, appears to be an exception; small males did not mimic females, most had their own nest and not once in 20 h of spawning observations did a second male enter a nest. In this field study, we ask if sneaker tactics are constrained and what factors favour independent nesting by small males. Larger males received more eggs sooner and attracted more females than smaller males. Larger males may be preferred because they released more sperm per ejaculation and fanned their eggs more often. In the long-term, however, larger males did not spawn more often than smaller males since: (1) the nests of large males filled up with eggs sooner than the nests of small males; (2) large males refused females more often than small males; and (3) a highly biased ratio of females to preferred large males meant many females had no option but to spawn with small males. Thus, small males nest independently, despite potential for female mimicry tactics (multiple females spawned simultaneously) and simultaneous parasitic spawning (males exited the nest during spawning)     

Brood defense and filial cannibalism in the spottail darter (<Emphasis Type="Italic">Etheostoma squamiceps)</Emphasis>: the effects of parental status and prior experience

Male spottail darters (Etheostoma squamiceps) defend nest sites in which females deposit eggs over the course of several weeks. In laboratory experiments, I tested three predictions of the hypothesis that the presence of eggs increases the value of a nest site to male spottail darters: (1) males should preferentially defend nests with eggs over empty nests, and males that sired the eggs should (2) spend more time defending them and (3) consume less of their broods than should alloparent males. Parent males were tested using eggs they were guarding in the field; alloparents included males that were not defending eggs immediately prior to the experiment (inexperienced), and males that were defending eggs in the field when captured but were given eggs sired by a different male (experienced). In choice experiments in which males were offered nest sites differing only in the presence of eggs, males preferred to defend nest sites with eggs regardless of parental status. Parent males spent significantly more time defending eggs and consumed less of their broods than did inexperienced alloparent males. Experienced alloparent males were similar to parent males in brood defense but had levels of filial cannibalism similar to inexperienced males, suggesting that prior experience may influence brood defense but not egg consumption.     

Female cannibalism and male courtship tactics in threespine sticklebacks

Summary Female threespine sticklebacks (Gasterosteus aculeatus) frequently raid male nests and eat all the eggs therein. We tested the hypothesis of Vickery et al. (1988) that females prefer to raid nests containing large numbers of eggs than ones with smaller numbers of eggs. This hypothesis is based on the finding that females spawning in nests containing many eggs will have reduced hatching success because of egg crowding. By consuming the male's eggs and forcing him to rebuild his nest, raiding females might obtain a new opportunity to spawn under better conditions. Our results were consistent with the first prediction of this hypothesis that females were more likely to spawn in nests containing fewer eggs than in nests with many eggs. However, this may be the result of males becoming less receptive to females as the number of eggs in their nests increases. Prediction 2 was that females should raid those nests containing the most eggs. Contrary to this prediction, males defending only one clutch were as likely to have their nests raided by groups of females as males defending several clutches of eggs. Female cannibalism is therefore unlikely to have evolved as a means of gaining access to a male defending a small number of eggs. We also examined the tactics used by males to counter female raids. Most raids occur when the male is courting, and nests are more vulnerable to shoals of females than to single females. Therefore, we hypothesized that males with eggs preferentially court a single female rather than large groups of females, and that males without eggs court both groups indiscriminately. We also predicted that males restrict the number of females they mate with when risk of having their nest raided is high. Our results indicate that: (1) both males with eggs and those without eggs minimize the risk of female cannibalism by courting solitary females rather than groups of females and (2) males limit the number of females that lay eggs in their nest when several potentially raiding females are present. Offprint requests to: G.J. FitzGerald     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Female spawning patterns and male mating success in the sand goby Pomatoschistus minutus

In June 1989 in a study conducted near Träminne Zoological Station, Finland (60° N 23° E) I investigated whether or not male mating success could be explained by female choice for male size in sand gobies (Pomatoschistus minutus). Male mating success was constrained by nest size and increased markedly with increasing nest size. I also found a negative correlation between the length of spawning females and the fullness of the nest. As large females lay more eggs, they also need to find a nest with more space available for the eggs. The size of males without eggs was the same across nest size, whereas the size of males with eggs significantly increased with increasing nest size. This is interpreted as female discrimination against males as mates in nests that are often contested. There was no correlation between a male's size and his mating success when males with no eggs in their nests were excluded. A male removal experiment, however, showed that, in a specific nest, when male size increases so does mating success, whereas, if male size decreases, mating success also decreases. It is concluded that in the sand goby females prefer to mate with larger males, especially when the male possesses a high-quality nest that he most probably will have to defend against other males.     

The artistic three-spined stickleback (<Emphasis Type="Italic">Gasterosteous aculeatus</Emphasis>)

We investigated whether female three-spined sticklebacks ( Gasterosteus aculeatus) are more attracted to a male whose nest contains colourful artificial material than to a nest that only contains algae. The purpose was to examine whether nest characters are of importance for female choice in the three-spined stickleback. Most studies on female preference in this species have focused on male morphological traits. In our study, on a marine population of three-spined stickleback on the West coast of Sweden, we found that males marked their nests' entrance with a deviant colour of algae — an apparent decoration. Moreover, males also decorated their nests with shiny and colourful foil sticks and spangles when provided with such, thereby apparently reducing the nest camouflage. In a mate choice experiment, females were more attracted to males with nests containing sticks and spangles than to males with undecorated nests. Nest decoration may be important for the evolution of paternal care in this species: if males advertise their paternal skills by ornamenting their nest they will receive more matings than males with dull nests.     

Shiny cowbirds share foster mothers but not true mothers in multiply parasitized mockingbird nests

Obligate brood parasitic birds, such as cowbirds, evade parental care duties by laying their eggs in the nests of other species. Cowbirds are assumed to avoid laying repeatedly in the same nest so as to prevent intrabrood competition between their offspring. However, because searching for host nests requires time and energy, laying more than one egg per nest might be favoured where hosts are large and can readily rear multiple parasites per brood. Such ‘repeat parasitism’ by females would have important consequences for parasite evolution because young parasites would then incur indirect fitness costs from behaving selfishly. We investigated shiny cowbird (Molothrus bonariensis) parasitism of a large host, the chalk-browed mockingbird (Mimus saturninus), in a population where over 70 % of the parasitized mockingbird nests receive multiple cowbird eggs. We assessed egg maternity directly, using cameras at nests to film the laying of individually-marked females. We also supplemented video data with evidence from egg morphology, after confirming that each female lays eggs of a consistent appearance. From 133 eggs laid, we found that less than 5 % were followed by the same female visiting the nest to lay again or to puncture eggs. Multiple eggs in mockingbird nests were instead the result of different females, with up to eight individuals parasitizing a single brood. Thus, while cowbird chicks regularly share mockingbird nests with conspecifics, these are unlikely to be their maternal siblings. Our results are consistent with shiny cowbird females following a one-egg-per-nest rule, even where hosts can rear multiple parasitic young.     

Why do males tolerate sneakers? Tests with the European bitterling, Rhodeus sericeus

In most species, males attack other males that attempt to gain fertilizations through sneak copulations. Here we report on a system where dominant males show a low level of aggression against sneakers at the initial stages of territory establishment. Females of the European bitterling, Rhodeus sericeus, lay their eggs in living mussels and males fertilize the eggs by releasing sperm over the mussels both before and after egg laying. When we allowed males to court females to a mussel containing no eggs at different male densities - one, two, four, or six males - the dominant male showed a low level of aggression against other males that released sperm. The dominant male became aggressive toward the other males only after eggs had been laid. This unusual pattern could be due to either some benefit of accepting sneakers or a high cost of aggression. We found support for both possibilities. The presence of several males decreased the time until a female spawned, whereas increased aggression by the dominant male against other males during a second female presentation, when the male was more territorial, interrupted courtship and increased the time until spawning. Females appeared to be attracted by both the presence of several males around a mussel and increased courtship under male competition. The bitterling mating system possibly differs from that of other species due to lack of investment in nest building and parental care, and high costs of defending the spawning site against sneakers.     

Multiple mate choice criteria and the importance of age for male mating success in the microhylid frog, Cophixalus ornatus

We examined multiple mate choice criteria in Cophixalus ornatus, a terrestrial breeding, microhylid frog. Mate choice consisted of three stages: mate attraction (male calling), courtship (male behavior between the call site and the nest), and nest site selection by the female. For male C. ornatus, the possession of a call with low dominant frequency relative to calling neighbors increased the probability that they would attract females. Dominant frequency was negatively correlated with age independent of male mass and snout vent length. When escorting the female from the call site to their nest, males traveled along more convoluted paths than when returning to the nest alone. The convolution of the path was, therefore, considered an aspect of courtship. Females released eggs into nests with structural characteristics typical of nests constructed by older males. Thus, females increased their chances of locating an acceptable nest by preferentially approaching males with lower dominant frequencies. This study is the first to demonstrate that age, independent of mass or snout-vent length, can influence call characteristics in anurans, and it is also the first to demonstrate the importance of male age to female mate choice in an amphibian.     

Female sand gobies gain direct benefits by choosing males with eggs in their nests

In some fish species with paternal care, females prefer to spawn with males whose nests already contain eggs. Several hypotheses have been put forward to explain this behaviour, such as reduced risk of predation or cannibalism (the dilution effect), increased parental investment, and mate copying. This experimental study focuses on female mate choice in the sand goby, Pomatoschistus minutus. Females were found to choose males with eggs in their nests. In addition, hatching success increased with clutch size, mainly because males with larger clutches showed less filial cannibalism. Increased egg survival in large clutches may thus be explained by a combination of the dilution effect and higher parental investment. In another experiment, females did not seem to copy the observed mate choice of other females. In conclusion, female preference for males with eggs in their nests is adaptive, and can be explained by direct benefits, as more surviving offspring are produced. Received: 23 December 1995/Accepted after revision: 11 May 1996     

Allopaternal care in the fathead minnow,Pimephales promelas: females prefer males with eggs

Summary Some species of fishes with exclusive male parental care exhibit the phenomenon of allopaternal care; that is, some males acquire and care for other males' eggs. We conducted a series of experiments to investigate the dynamics and evolution of allopaternal care in one such species, the fathead minnow (Pimephales promelas). In choosing a nest site, a newly reproductive male tended to take over the nest site of a parental male by evicting the resident male, rather than occupy a physically identical empty nest site. The new male generally cared for the old male's eggs, and in most cases, daily egg survival improved under the new male's care. When males were given a choice among unguarded nest sites, they preferred to occupy nest sites already containing eggs. When eggs were randomly assigned to nesting males, females preferred to spawn with males who had eggs in their nest sites. Thus, it appears that female preference for males with eggs led to the evolution of allopaternal care in the fathead minnow.     

Aggregation and dispersal based on social cues as a nest-site selection strategy in a resource-defence polygynandry mating system

Many animals must choose a nest site in order to reproduce. However, it is unclear how nest-site selection strategies vary across different mating systems. We must therefore explore nest-site selection strategies in a range of mating systems, including the interaction between resource-defence polygyny and polyandry (i.e. polygynandry). In this study, we imposed a re-settlement event in the terrestrial toadlet Pseudophryne bibronii and measured the influence of the spatial position of each male’s nest site with respect to rival males on the likelihood that it would be abandoned or receive eggs. We captured every calling male in a population, measured their breeding success and released them back into the breeding area. We then recorded the establishment and abandonment of nest sites by males over 26 consecutive nights. Spatial positioning did not have any significant effects on male-breeding success, supporting claims that females show less discrimination between nest sites when they are polyandrous and spread their eggs amongst multiple male nests. However, we found that males consistently selected nest sites according to a site’s spatial position, which suggests that fitness benefits unrelated to male breeding success (e.g. reduced mortality risk) might influence male nesting decisions. Overall, our study provides new evidence that the mating system adopted by a population can influence the cues that individuals respond to when selecting nest sites.     

Egg mimicry and allopaternal care: two mate-attracting tactics by which nesting striped darter (Etheostoma virgatum) males enhance reproductive success

In a variety of fish species with paternal care of offspring, females prefer to spawn in nests that already contain eggs. This female preference has been hypothesized to explain egg thievery in male sticklebacks, allopaternal care of eggs in minnows, and the evolution of egg-mimicking body features in male cichlids and darters. Here we employ microsatellite-based parentage analyses to evaluate the reproductive success of striped darter (Etheostoma virgatum) males that appear to utilize two of these functionally related tactics to entice females to spawn in their nests. In an isolated population (Clear Creek, Ky.), we observed that breeding males develop conspicuous white spots on their pectoral fins. If these spots are egg mimics, as we suspect, then this represents the fourth independent evolutionary origin of egg mimicry documented to date in darters, the first based on pigmentation (as opposed to physical structures), and the first in which the egg mimics vary greatly in number among males. From direct counts of microsatellite genotypes in clutches of embryos, at least 3.8 females contributed to the progeny within a typical nest, and females tended to spawn preferentially with males that were larger and displayed more egg-mimic spots. In another population (Hurricane Creek, Tenn.) without egg mimics, the multi-locus genetic data document that allopaternal care is common, especially among the smallest males who sometimes tend nests containing their own as well as an earlier sire's offspring. Thus, these foster males had adopted egg-containing nests and then successfully spawned with subsequent females. Overall, the genetic data on paternity and maternity, in conjunction with field observations, suggest that egg mimicry and allopaternal care are two mate-attracting reproductive tactics employed by striped darter males to exploit female preferences for spawning in nests with 'eggs'. Electronic Publication     

Mate guarding in the swallowHirundo rustica

Summary The importance of mate guarding by males in the monogamous swallowHirundo rustica was studied by temporarily detaining the males. Mate guarding reduced the frequency of extra-pair copulations and of sexual chases involving female mates. Males participated in sexual chases more frequently if they had a non-fertile female. Neighbouring males of ‘widowed’ females increased their own mate guarding presumably in response to the experimentally increased rate of sexual chases. Neighbouring males with a fertile female increased their mate guarding more than did males with a non-fertile female. Addition of eggs to swallow nests in the post-fledging period of the first brood induced mate guarding by male nest owners. These males also copulated more frequently with their mates than did control males. Neighbouring male swallows responded to the increased mate guarding by showing sexual interest in the guarded females. removal of eggs from swallow nests during the laying period, leaving only one egg in the nest, resulted in reduced nest attendance by females. Male mates responded by increasing their mate guarding intensity as compared to controls, and neighbouring males showed an increased sexual interest in these females.     

Extrapair copulations in western bluebirds (Sialia mexicana): female receptivity favors older males

. Extrapair paternity has been the focus of numerous studies of socially monogamous birds, but the dynamics of extrapair male-female interactions have been explored in relatively few. In western bluebirds (Sialia mexicana), a species with 20% extrapair paternity, two to four observers monitored extrapair copulations (EPCs) occurring on the female's home territory while her social mate was detained in a cage near the nest. Detention elevated the frequency and visibility of EPCs, so we could score receptivity of females to identifiable extrapair males without the interference of intermale aggression. Most intruding extrapair males were neighboring breeders visiting outside their own mate's fertile period. Local male relatives of the detained male attempted nonincestuous EPCs, but neighboring sons did not attempt to copulate with their mothers and, in some cases, chased away intruders. Female receptivity was not influenced by whether the detained mate was visually present or hidden behind cloth and did not increase with the persistence or relative size of the extrapair male. Female receptivity increased with the age difference between the extrapair male and the female's social mate, suggesting that females prefer older males as extrapair partners. This study suggests that costs of resistance are relatively unimportant in determining female receptivity in western bluebirds, but identifies preferential receptivity to older males as a potential factor in the outcome of inter- and intrasexual conflict over paternity.     

Are nest characters of importance when choosing a male in the fifteen-spined stickleback (Spinachia spinachia)?

Male fifteen-spined sticklebacks (Spinachia spinachia) build their nests by wrapping epiphytic algae around macroalgae and securing them with secretional threads. In this study, I show that female fifteen-spined stickleback prefer males which have built their nests above the surrounding vegetation. High-located nests are safer, because they are less likely to attract egg-predators such as shore crabs (Carcinus maenas). Thus, I found that shore crabs more rapidly find eggs in nests built close to the bottom than in nests well above the it. Moreover, male-male competition could be an additional explanation as to why males build nests high up. Thus, my field results suggest that the closer the males were to a neighbour, the higher they built. Larger males were also found to have larger territories but male size did not correlate with the height of the nest above the surrounding vegetation. Females showed no preference for nest size, regardless of the presence of egg-predators. However, they tended to choose nests that had no egg-predators in the vicinity over nests with egg-predators close by. Female choosiness for certain nest characters may provide one explanation for the evolution of male care in this species. Received: 29 December / 1999 Revised: 18 April 2000 / Accepted: 28 May 2000     

Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

Sexual dimorphism and male mating success in the tentacled blenny, <Emphasis Type="Italic">Parablennius tentacularis</Emphasis> (Teleostei: Blenniidae)

Although external sexually dimorphic traits are commonly found in males of combtooth blenny species, little is known about the benefit they can convey to male mating success. Indeed, while female preferences for large males have been demonstrated in some species, the possible role played by dimorphic ornaments has been neglected. We now report on the tentacled blenny, Parablennius tentacularis, a species where males are characterized by bulb glands on the anal fin and both sexes exhibit a dark spot on the dorsal fin and orbital tentacles. Males are territorial, make nests in empty bivalve shells, and provide solitary parental care for the eggs. Using morphometric analysis and field collected data on male and female external features, nest characteristics and number of eggs in the nests, we have assessed the development of dimorphic traits in both sexes and male mating success. The results reveal that orbital tentacles of males are more developed and more variable in size than those of females. Larger males exhibit longer orbital tentacles and larger anal glands but do not necessarily occupy larger nests. Male mating success is significantly correlated with the inner nest surface area and with orbital tentacle size but not with body size. These results provide support for a primary role of male ornaments in enhancing blenny male mating success and are discussed in the context of mate choice for direct and indirect benefits.     

Female control of extra-pair fertilization in tree swallows

Summary In a Canadian population of tree swallows, Tachycineta bicolor, DNA fingerprinting has previously shown that half of all broods contain many offspring resulting from extra-pair copulations (EPCs), whereas the other half contain only legitimate offspring. This bimodal pattern of extra-pair paternity might be due to variation in the effectiveness of male paternity guards, variation in female ability to resist EPCs, and/or variation in female pursuit of EPCs. Here we report experimental evidence for female control of copulations and fertilizations and the occurrence of two alternative copulation strategies among females in this population. Ten paired male tree swallows were removed on the day their mates laid the first egg. Replacement males took over the nestbox within 0.5–23 h and attempted to copulate with the widowed female. Assuming that eggs were fertilized approximately 24 h prior to laying, the first two eggs were fertilized before the male was removed, while the third and subsequent eggs could potentially be fertilized by the replacement male. Fingerprinting revealed that the first two eggs were sired by the resident males in five nests and by extra-pair males in the remaining five nests. The widows that had been faithful to their initially chosen mate rejected copulation attempts by the replacement male until most of the eggs had been laid. Consequently, nearly all eggs laid by these females were sired by the original male. The widows that had been unfaithful prior to male removal copulated sooner with the replacement male than females that were faithful to their mate. However, these replacement males also had a very low fertilization success; most eggs were sired by males that were not associated with the nest. This is consistent with the situation in non-experimental nests where unfaithful females copulate with their mate at the same rate as faithful females, yet unfaithful females have a majority of offspring sired by extra-pair males. We conclude that fertilization patterns to a large extent are determined by the female through active selection and rejection of copulation partners, though our results also allow some speculation that females have control over sperm competition. Female copulation tactics are probably determined some currently unknown fitness benefits of having the offspring sired by particular males.Correspondence to: Raleigh J. Robertson     

Cues used by shiny cowbirds (Molothrus bonariensis) to locate and parasitise chalk-browed mockingbird (Mimus saturninus) nests

Unlike other birds, shiny cowbirds (Molothrus bonariensis) must locate host nests where to lay their eggs and then decide whether to parasitise them. They should also synchronise their laying with that of the host to increase the survival of parasite egg and young. Shiny cowbirds can discover nests using host behaviour as a cue, or by searching the habitat without need for the presence of a host. Besides, they can synchronise parasitism with host laying by monitoring nests during building and laying, or directly by assessing the degree of development of embryos through the puncture of host eggs. Alternatively, synchronization can arise by lower nest attentiveness during host laying. We determined the extent of synchronization between laying of shiny cowbirds and chalk-browed mockingbirds (Mimus saturninus) and estimated if parasitism was negatively associated with host nest attentiveness. We also conducted an experiment to test if host activity was necessary to locate nests, and if puncture of host eggs was a cue for deciding parasitism. Shiny cowbirds synchronised parasitism with host laying in 75% of the cases and synchronization was not explained by lower host nest attentiveness during laying. Shiny cowbirds located nests without need for presence of a host, but the decision of parasitising the nest depended on host activity at the nest. The information that shiny cowbirds could obtain through egg punctures was not necessary for deciding parasitism. Our results indicate that shiny cowbirds rely on the precise timing of their eggs and avoid laying in unsuitable nests.