Effect of clutch size on male egg-fanning behavior and hatching success in the goby, Eviota prasina (Klunzinger)

In fish that exhibit paternal care, the females often choose their mates on the basis of male traits that are indicative of the parental ability of the males. In a marine goby, Eviota prasina, males tend their eggs within their nests until hatching, and females prefer males that have longer dorsal fins and exhibit courtship behavior with a higher frequency as their mates. In order to clarify the relationship between these sexually selected traits and the parental ability of males of E. prasina, the factors affecting the hatching success of eggs within male nests and the male parental care behavior were examined in an aquarium experiment. Females spawned their eggs in male nests and the clutch size of females showed a high individual variation (range = 88-833 eggs). The hatching success of eggs within male nests showed a positive correlation with the time spent by males in fanning eggs and the clutch size. In contrast to the prediction, however, the hatching success did not show a significant correlation with the sexually selected traits, i.e., the male dorsal fin length and the frequency of courtship displays. Moreover, multiple regression analysis indicated that the time spent by the males in fanning was the most important factor affecting the survival rate of the eggs. The time spent by males in fanning behavior was influenced by the clutch size within their nests; the fanning behavior of males occurred with a higher frequency when they tended larger clutches. Males are required to invest a greater effort in egg-tending behavior to achieve a higher hatching success when they receive larger clutches, probably due to the greater reward for their parental behavior. Based on their mate choice, females may obtain other benefits such as high quality offspring.     

Nocturnal hatching timing of mouthbrooding male cardinalfish Apogon niger

We observed hatching behavior by mouthbrooding males of the cardinalfish, Apogon niger. Mouthbrooding males showed no feeding activities at night, in spite of their nocturnal feeding habit. On the day of hatching, they released newly hatched larvae from their mouths on average 81 min after sunset. Semilunar hatching periodicity was significant, but its diel pattern was independent of the tidal rhythm. Sunset hatching may be advantageous not only to offspring because of their low predation risk but also to parental males because they can resume feeding sooner, thereby reducing the energetic loss from fasting while mouthbrooding. Received: August 22, 2000 / Revised: November 28, 2000 / Accepted: January 12, 2001     

Courtship feeding in Ospreys Pandion haliaetus: a criterion for mate assessment?

In many bird species males provision their mates prior to egg-laying. Courtship feeding has been suggested to function in several ways: to advance laying date by improving female condition, to induce a female to copulate or to allow a female to assess her mate. The role of courtship feeding in Ospreys Pandion haliaetus was investigated in British Columbia, Canada. Courtship feeding rate affected the probability of a pair initiating a clutch. Pairs that laid eggs had higher rates of courtship feeding than pairs that did not lay eggs in both 1991 and 1992. Male courtship feeding rate also correlated negatively with the duration of the courtship period. Experimentally increasing the amount of food available to females prior to egg-laying resulted in a nonsignificant reduction in the duration of the courtship period. This study found no evidence to support the suggestion that female Ospreys trade copulations for food during the courtship period; only 63 of 385 copulations observed were associated with feeds, and courtship feeding rate did not correlate with the copulation rate of a pair. Male provisioning rates, however, were predictable; courtship feeding rate correlated with both male delivery rate to the nest when chicks were 1–2 weeks old and mean brood growth rate. Female Ospreys therefore may be able to predict the quality of subsequent paternal care using courtship feeding rate. As predicted if optimal hatching asynchrony is dependent on food availability, mean brood growth rate, an indirect measure of male parental care, was negatively correlated with hatching asynchrony. This suggests that female Ospreys may manipulate hatching asynchrony in response to male courtship feeding rate, thereby maximizing the productivity of their brood at predicted food levels.     

Unravelling the evolution of complex reproductive traits with phenotypic engineering

The social dynamics surrounding courtship, mating and parental care are complex enough when just a single male and female are involved, but for species that employ multiple strategies for achieving fertilization success, the network of interactions among rivals, allies and suitors can be utterly complicated. Such is the case in the ocellated wrasse, Symphodus ocellatus, in which males adopt one of three mating strategies. The large, colourful “nesting males” court females, defend territories and care for fertilized eggs until they hatch. The smaller “satellite males” help the nesting males court females and guard against the third morph, the “sneaker males”, which sneak in when a nesting male is spawning with a female and surreptitiously release sperm. Sneaker males perform no courtship displays nor defend territories, so their reproductive investment is devoted entirely to sperm production. And these alternative male strategies work: 100% of nests contain some eggs fertilized by sneaker and satellite males, despite the fact that parental care is solely the responsibility of nesting males In this issue of Molecular Ecology, work to untangle the proximate mechanisms regulating the reproductive physiology of nesting males and their behaviour towards other males, which impacts the entire social network. Moreover, they describe how variation in neuroendocrine regulation can give rise to variation in reproductive traits, upon which sexual selection can act.     

Costs of egg ventilation for male common gobies breeding in conditions of low dissolved oxygen

This study examines the effects of harsh environmental conditions on life history trade-offs in parental care in a marine fish, the common goby, Pomatoschistus microps. We compared male parental care and hatching success over two sequential brood cycles in fish breeding in conditions of either low dissolved oxygen or normal levels of oxygen. Males compensated for a low oxygen environment by increasing the time they spent fanning water over their eggs, as well as their fanning tempo. They also increased the frequency of egg-directed activities and decreased nest-building activities. Males in the low oxygen treatment lost more weight than control fish during the first spawning, and were more likely to abandon care during the second spawning. Males that completed care under low oxygen conditions did not differ from control males in the hatching success of their offspring or the size of young at hatching, but hatching started on average 1 day later. Thus, greater parental allocations to offspring while breeding in a harsh environment led to reduced future allocations. Copyright 1999 The Association for the Study of Animal Behaviour.     

Dietary carotenoid availability influences a male's ability to provide parental care

Despite convincing evidence that carotenoid availability canhave positive physiological effects, we still lack informationon the functional consequences of carotenoid limitation at thebehavioral level. Given the role carotenoids play in mitigatingoxidative stress produced during physical activity and as immunostimulants,one behavioral function on which they may have a significantimpact is an individual's capacity to provide parental care.We tested this hypothesis using three-spined sticklebacks (Gasterosteusaculeatus), a species in which males provide obligate and intensivepaternal care. Males were fed either high or low (but biologicallyrealistic) levels of carotenoids and monitored throughout incubation,during which we quantified 2 key aspects of parental care: theirability to fan their eggs under normoxic and hypoxic conditions(when both the costs and requirements of fanning increase) andtheir ability to defend their nest against a simulated conspecificmale. High-carotenoid diet males fanned their eggs at a significantlyhigher rate during hypoxic (but not normoxic) conditions andhad higher clutch hatching success than males fed the low-carotenoiddiet. There was no evidence that they defended their nest moreaggressively. Furthermore, low-carotenoid diet males also appearedto engage in cannibalization of their clutch. These resultsdemonstrate that dietary carotenoid availability can affecta male's ability to provide parental care, and we discuss thepotential mechanisms and implications of this finding.     

Courtship as an honest indicator of male parental quality in the bicolor damselfish, Stegastes partitus

Previous work on the bicolor damselfish, a species with exclusivemale parental care of eggs, suggested that female mate choicewas based on male characteristics. The aims of this study wereto determine whether females discriminate among potential mateson the basis of courtship and, if so, to determine whether courtshipserves as an indicator of male parental quality. Observationsmade over two reproductive cycles showed that courtship ratesand mating success of individual males are positively correlatedand that males begin courting several days before females beginlaying eggs. Experimental manipulations showed that a male'scourtship rate is indicative of the subsequent egg survivalfrom his nest. We suggest that observed differences betweenmales in their courtship rates and parental ability may be aresult of differences in their energy reserves. These resultsdemonstrate the operation of honest advertising and lend supportto adaptive models of sexual selection. [Behav Ecol 1991;2:295–300]     

Condition‐dependent male copulatory courtship and its benefits for females

Postcopulatory sexual selection has shaped the ornaments used during copulatory courtship. However, we know relatively little about whether these courtship ornaments are costly to produce or whether they provide indirect benefits to females. We used the mealworm beetle, Tenebrio molitor, to explore this. We challenged males using an entomopathogenic fungus and compared their courtship (frequency of leg and antennal contacts to the female), copulation duration, number of eggs laid, and hatching rate against control males. Infected males copulated for longer yet they reduced their leg and antennal contacts compared to control males. However, there was no obvious relation between infection, copulation duration, and courtship with egg production and hatching success. In general, our results indicate that the ornaments used during postcopulatory courtship are condition‐dependent. Moreover, such condition dependence cannot be linked to male fitness.     

Androgen and behavior in the male three-spined stickleback,Gasterosteus aculeatus. II. Castration and 11-ketoandrostenedione effects on courtship and parental care during the nesting cycle

Courtship declines and ceases while parental care increases in the presence of developing eggs during the nesting cycle of the male three-spined stickleback, Gasterosteus aculeatus. Furthermore, circulating 11-ketotestosterone (11KT) levels are higher during the initial "courtship phase" than during the later "parental phase," similar to that found in other paternal fishes. This study aimed to investigate a possible functional relationship between changes in 11KT levels and changes in reproductive behavior during the nesting cycle. To this end, groups of nonspawned and spawned male sticklebacks were sham-operated, castrated, or castrated and treated with 11-ketoandrostenedione (11KA), and the effects of the treatments on courtship and parental care were studied. Castration removed circulating 11KT, while 11KA replacement prevented the natural decline in 11KT during the parental phase (11KA converts to 11KT extratesticularly), as assessed by radioimmunoassay. Regardless of treatment, parental care remained low and courtship was present in all nonspawned males, even at the end of the experiment. However, courtship did eventually decline in castrated nonspawned males compared to the other two nonspawned groups. In all treatments of spawned males there was a drastic decline in courtship and an increase in parental care. In castrated spawned males, however, the decline in courtship came earlier than in the other two spawned groups. 11KA treatment did not prevent the natural decline in courtship/increase in parental care in spawned males, indicating that the natural decline in 11KT is not responsible for the main portion of the rapid changes in these behaviors over the stickleback's nesting cycle. The limited effects of castration also exclude other gonadal hormones from being responsible for most of these changes.     

Predominant maternal egg care and promiscuous mating system in the Japanese filefish,Rudarius ercodes (Monacanthidae)

Synopsis Reproductive behavior of the Japanese filefish, Rudarius ercodes, was studied at the rocky reef off Koinoura, northern Kyushu, Japan, between June and October 1989. Aggressive display was observed between males, but they were not territorial. Males had four types of courtship behavior: vibrating, tail bending, leaning and nuzzle. Spawning occurred early in the morning. A female and 1–3 male(s) mated together on brown algae. Each female spawned repeatedly with an interval of 6–12 days. Females cared for eggs and embryos from just after spawning until hatching, 2–4 days. Female egg care consisted of tending and guarding. Females tended eggs by blowing water on them and by fanning them with their pectoral fins. Females guarded eggs by driving away fish passing nearby. In some cases, males also guarded eggs by staying near the eggs and driving away conspecific males. Whether a male cares for eggs with a female seems to be affected by the form of mating (pair mating or single female-multiple male mating), and the probability of further reproduction after spawning. Dominant males showed a tendency to pair with a specific female intermittently over a two-month period. Mating, however, did not always occur between members of such pairs, and mates appeared to be inter-changeable with a promiscuous mating system.     

Steroid hormones and paternal care in the plainfin midshipman fish (Porichthys notatus)

The present study investigated the relationship between plasma steroid hormone levels and the expression ofpaternal behavior in the plainfin midshipman fish (Porichthys notatus), where males may simultaneously care for multiple clutches in different stages of development. Blood samples were collected from free-living parental males during that part of the breeding season when males may be found in various stages of parental care. Plasma 11-ketotestosterone levels were significantly higher in males with empty nests and nests containing only eggs than in males with nests containing embryos. All males with nests containing embryos had undetectable testosterone levels, whereas testosterone levels were detectable in many males with empty nests or nests containing only eggs. Estradiol levels were detectable in only a few males from nests with no eggs or nests containing only eggs. Cortisol levels were not correlated with stage of paternal care or with handling time. These results follow the frequently reported vertebrate pattern of declining androgen levels over the course of the breeding season or during the period of parental care. However, many male midshipman guarding nests containing only eggs had androgen levels similar to those of males whose nests contained no offspring. Thus the pattern of androgen levels exhibited by reproductively active parental male midshipman may reflect a compromise between investment in paternal care versus courtship and/or territoriality.     

The Influence of Parental Status on Courtship Effort in a Paternal Caring Fish

It is widely assumed that caring for young limits the motivation of parents to seek additional mating opportunities. However, in situations where parental care does not involve direct provisioning of the offspring, but rather activities directed at the brood as a whole (e.g. guarding), it may be more efficient for parents to care for large numbers of young at once. This may be especially true for species with exclusive paternal care, with fathers that have recently acquired a brood of young potentially benefitting from vigorously courting prospective mates, so as to maximise their chances of attaining a large number of young to rear together. We experimentally tested this hypothesis in the three‐spined stickleback (Gasterosteus aculeatus), a fish with male only care. Contrary to our predictions, we found no evidence of any differences in courtship between recently spawned egg‐tending fathers and males that had not spawned. However, males that were permitted to spawn, but then had their eggs taken from them, courted less vigorously. Together, the results of our study suggest that the potential benefits of vigorous courtship in terms of acquiring additional young may be offset by additional costs faced by parental males.     

Evolution of Female Egg Care in Haremic Triggerfish,Rhinecanthus aculeatus

Paternal care is predominant among telcost fishes with external fertilization. This study describes maternal care in a haremic coral-reef fish and discusses the possible factors leading to its evolution. Both sexes of the triggerfish Rhinecanthus aculeatus (Balistidae) maintained territories; some individuals for more than 8 years. Each male's territory overlapped 2-3 female territories. Pair-spawning occurred around sunrise. Only females cared for the demersal eggs until hatching, which occurred just after sunset on the day of the spawning. No predation was observed on eggs under the maternal care, but experimental removal of parental females decreased the hatching rate to nearly zero. Egg-guarding females foraged as frequently as males, but less than half of non-spawning days. Spawning occurred only in the periods of about 1 wk around the new or full moon, and individual females spawned up to three times in each period. Thus, the maternal care did not significantly affect the duration of the females' spawning intervals, while males would suffer mate loss if they performed parental care. In this situation, maternal care should be the evolutionarily stable strategy. Evolutionary transition from no care to maternal care and then to biparental care is suggested in the Balistidae.     

Male parental care and its adaptive significance in a neotropical frog

In the Puerto Rican frog Eleutherodactylus coqui, parental care is performed exclusively by males, and consists of attending the eggs and hatchlings at a terrestrial oviposition site. The two major behavioural components of parental care are egg brooding and nest defence against conspecific egg cannibals. Defence behaviour includes aggressive calling, biting, sustained biting, wrestling, and blocking directed against nest intruders. Parental care lasts from oviposition to hatching (17–26 days) and often for several days after hatching. During pre-hatching development, males are present in their nests 97.4% of the time during the day and 75.8% of the time at night. A large portion of this time is spent brooding eggs. In a field experiment, males were removed from their nests and the fate of clutches was monitored. Compared to control clutches (males not removed), experimental clutches had significantly lower hatching success and suffered significantly greater mortality from desiccation and cannibalism. Hence, parental care yields significant benefits to male fitness via increased offspring survival.     

Parental quality and egg size affect chick survival in a precocial bird, the lapwing Vanellus vanellus

Effects of egg size and parental quality on lapwing Vanellus vanellus chick survival were studied in southwestern Sweden over 6 years. Chicks from large eggs were heavier at hatching and survived significantly better than those from small eggs. To control for the confounding effect of parental quality on egg size and chick survival, we performed a cross-fostering experiment during 2 years, exchanging clutches between nests with large and small eggs. In control clutches, chicks from large eggs survived better than those from small eggs, but we found no significant difference in chick survival between exchanged clutches. Thus, egg size did not affect chick survival independently of parental quality. Fledging success increased with parental age and/or experience, and with female body mass. Hence, both egg size and parental quality affect chick survival in the lapwing. Received: 22 February 1996 / Accepted: 30 September 1996     

Spawning behavior of the pygmy leatherjacket Brachaluteres jacksonianus (Monacanthidae) in southeastern Australia

Spawning behavior of the pygmy leatherjacket Brachaluteres jacksonianus was studied in southeastern Australia. Prespawning females thrust the snout repeatedly into spawning substrata, while males exhibited courtship display and nuzzled the female’s face. The females and males touched their abdomens in pairs and released gametes. No parental egg care was observed after spawning. Fertilized eggs measured 0.74 mm in diameter, and each clutch contained 50–191 eggs. The eggs were attached to filamentous red algae, which were found on the surface of the seagrass and other substrata. Reproductive ecology of B. jacksonianus is similar to that of congeneric Brachaluteres ulvarum, although selection of spawning substrata differed in two species.Supplementary material to this paper is available in electronic format at http://dx.doi.org/10.1007/s10228-005-0272-8     

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     

Parental fish change their cannibalistic behaviour in response to the cost-to-benefit ratio of parental care

Partial filial cannibalism, the act of cannibalizing some offspring, has been explained as a response to the high energetic cost of care. I tested this hypothesis by manipulating the cost-to-benefit ratio of care in the scissortail sergeant, Abudefduf sexfasciatus, a tropical damselfish with male care. Background egg mortality was lower than the incidence of cannibalism, confirming that males did not just dispose of dead eggs. Investment in the current brood affected future investment, because males forced to skip a brood cycle put more effort into courtship during the following cycle and obtained larger broods than did unmanipulated males. Any factor influencing the cost-to-benefit ratio of parental care should also affect the incidence of cannibalism. I reduced the cost of care by supplementary feeding and reduced the benefit of care by simulating a decrease in paternity certainty through simulated intrusions by non-nesting males. Supplementary feeding significantly reduced partial filial cannibalism by parental males, a result compatible with the hypothesis that eggs are consumed to cover the energetic costs of parental care. Cannibalism decreased regardless of whether males were fed with conspecific eggs or crabmeat. Cannibalism was only reduced but not fully eliminated by supplementary feeding, and residual levels of cannibalism after feeding were similar to the background rate of egg mortality. Simulated intrusions by non-nesting males led to an increase in filial cannibalism and a decrease in parental effort.     

Mate availability affects female choice in a fish with paternal care: female counterstrategies against male filial cannibalism

Theory predicts that individuals should adopt counterstrategies against intersexual conflict with their mating partners if the counterstrategies are effective and cost-efficient. In fishes, males with parental care often cannibalize their own offspring, which reduces the female’s fitness and creates intersexual conflicts. Males of the goby Rhinogobius flumineus cannibalize more eggs in the nest when they have access to additional females prior to spawning. Thus, it is predicted that females will strategically avoid spawning with males that have high mate availability. In the present study, we experimentally tested this prediction. When sexual pairs were placed in tanks, most females (control females; 21/22) successfully spawned inside the nest. In contrast, when a gravid female (stimulus female) that was housed in a small transparent cage was shown to the experiment pairs prior to spawning, only about half of the females (experiment females; 16/29) spawned inside the nest; the remaining females released unfertilized eggs outside of the nest. Moreover, experiment females infrequently accepted and followed males into nests, and delayed spawning more often than control females. R. flumineus females prefer males that court frequently. Indeed, experiment females that infrequently received courtship tended to spawn outside of the nest. However, infrequent courtship alone could not explain outside-nest spawning, delay in spawning, or the shorter stay of females in nests. These results imply that the presence of a stimulus female dampens female spawning with males. We suggest that R. flumineus females may strategically reject or hesitate to spawn with males that have high mate availability, and that this spawning avoidance may be a counterstrategy against male filial cannibalism.     

Honesty of agonistic signalling and effects of size and motivation asymmetry in contests

Game theoretical models predict that the main function of fighting behaviour is to assess the relative fighting ability of opponents. The sequential assessment game has often been used to investigate contests, while honest signalling theory has received much less attention. With the wolf spider Hygrolycosa rubrofasciata we investigated whether male agonistic signalling can reveal honest information about fighting ability, and how size and motivation asymmetries affect male fighting behaviour. We also determined whether male–male competition affects the courtship behaviour of the males. We found that agonistic drumming activity is an honest indicator of male fighting ability, and that relative size asymmetry and motivation to fight both contribute to the fighting ability. We also found that male–male competition decreases the courtship drumming rate of subdominant males, suggesting that male–male competition limits the opportunities for female choice. We conclude that sequential assessment and honest signalling may both be used in settling contests, and that they may be used simultaneously. Received: 10 December 1998 / Received in revised form: 23 June 1999 / Accepted: 5 July 1999