Plasticity in spawning visits of female damselfish, Stegastes nigricans : effect of distance to mates

Both sexes of the damselfish Stegastes nigricans hold individual territories in which they feed on filamentous algae. At dawn, females visit males' territories to spawn, and the males guard the eggs until hatching. We examined how females' spawning behavior varied according to the distances to their mates. Females usually mated with a single male per morning. The distance to the territory of a mate (0.7–12.8 m) did not affect a female's total spawning time per morning, but affected the number of her spawning visits with that male. Females made many repeated spawning visits when spawning with males at short distances, while they spawned the entire clutch in one visit when spawning with males at long distances. This plasticity in female behavior appears to be related to two costs during the spawning visits: (1) intrusions by other fish to feed on algae in the female's territory during her absence, which may cause the female to return repeatedly to her territory for defense, and (2) attacks on the female by other territorial fish, which increased with the distance to their mates' territories. To minimize the sum of both costs, females should change the number of spawning visits depending on the distance to the males' territories. Received: 30 September 1996 / Accepted after revision: 17 March 1997     

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     

Patterns of the electric organ discharge during courtship and spawning in the mormyrid fish,Pollimyrus isidori

Summary Pollimyrus isidori's electric organ discharge (EOD) is of the pulse type. Patterns of EOD intervals were investigated prior to, during and following spawning behaviors as related with overt behaviors, and with the sound production by the nestbuilding male. Prior to the time of reproduction, isolated and socially interacting fish (n=15) showed characteristic discharge interval patterns for resting, swimming, probing, hovering and hiding activities. Males (n=8) and females (n=6) did not differ in their mean EOD repetition rates during resting (11.6±2.5 Hz), nor Short Bursts/min (less than 20 intervals of 8–13 ms). In interacting fish Long Bursts (greater than 20 intervals of 8–13 ms, lasting for more than 300 ms) were observed only during the attack and bite sequence. A pursuing fish displayed a rapid alternation of Long Bursts with Discharge Breaks (300–1000 ms silence) during the chase behavior. Avoidance behavior which followed from several attacks was correlated with a Medium Uniform Rate (8–12 Hz) normally lasting for 20 to 60 s, or a Discharge Arrest (silence greater than 1 s) in the submissive fish. The nocturnal courtship behavior began soon after dark (1900 h). Spawning typically started 2 to 5 h after dark, continuing for 2 to 6 h until about 0200 h. During courtship and spawning the female's brief visits (15–25 s) to the male's territory recurred every 30–60 s. At all other times the female was aggressively excluded from the nest region. Courtship and spawning behaviors are described along with the electrical displays identified from 19 spawnings in three fish pairs (from a total of 37 spawnings in 4 males and 4 females). Just prior to the onset of courtship behavior, with male territorial aggression beginning to decline, females switched from a Medium Sporadic Rate pattern (resting and hiding patterns; 13 Hz) to a Medium Uniform Rate pattern (6–8 Hz) while still in their hiding area. Females continued to display this uniform rate throughout the courship and spawning period, including the courtship and spawning bouts when Discharge Breaks or Arrests also occurred. This persistance distinguishes the courtship pattern from the similar avoidance pattern (see above). The male courtship and spawning EOD pattern was similar to the female's and unique for a territorial male. He switched from a High Sporadic Rate (swimming EOD pattern; about 18 Hz) to a regularized Medium Uniform Rate (about 9 to 11 Hz) only during courtship and spawning bouts, including 1–3 EOD Breaks during Vent-to-Vent coupling (average interval: 272±71 ms, n=37). No sooner had the female left the spawning site than he resumed displaying a High Sporadic Rate. This temporal correlation of reproductive behaviors with electrical displays suggests their instrumental role in mutual acceptance of mates. Males showed their sex-specific type of EOD phase-locking, the Preferred Latency Response, only during the first few hours of entry of a fish in their tank. Two females with EOD waveform features more typical of males also spawned repeatedly; waveform does not appear to be critical. Males stopped their nocturnal sound production for the later part of courtship and the whole spawning period. Except for infrequent attacks on the female between spawning bouts, the male did not resume singing until the end of spawning when all eggs were shed (around 0200 h); from this time on the male sang until dawn. The sequencing of the three acoustic elements (moans, grunts, growls) are described. A catalogue of discharge patterns correlated with overt behaviors (Tables 1, 2), and an integrated summary time table of P. isidori's complex reproductive behavior are presented.     

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.     

Effect of female molt stage and sex ratio on courtship behavior of the blue crab Callinectes sapidus

In many species, males and females actively participate in courtship, and the outcome of pre-mating interactions influences the mating success of both sexes. Female blue crabs, Callinectes sapidus, mate soon after their final molt to maturity; thus female molt stage dictates the timing of mating. In a field experiment, we manipulated female molt stage and sex ratio to test their effects on the courtship behavior of both sexes, if female behavior influences the behavior and pairing success of males, and if male courtship influences male pairing-success. Early-molt-stage females avoided males during courtship, whereas late-molt-stage females sought out males. As a result, males had to pursue and capture early-molt-stage females whereas males displayed to late-molt-stage females and more easily physically controlled them. Males sometimes abandoned late-molt-stage females, but this occurred more often when females were abundant. The rate at which females avoided males was positively correlated with that of males abandoning females, and males that were unsuccessful at pairing met with higher rates of female resistance than successful males, suggesting that female behavior influences male pairing-success. Unlike unsuccessful males, successful males more often made the transition between display and maintaining physical control of the female. At high male sex ratios, males initiated courtship more readily; thus both sexual competition and female behavior influence male courtship in this species. Received: 7 July 1996 / Accepted: 10 January 1998     

To court or not to court: reproductive decisions by male fiddler crabs in response to fluctuating food availability

For males, courting and foraging are often behavioral alternatives, which take time and consume energy. When males have a possibility of mating with receptive females, there may be a behavioral trade-off between courtship and feeding; the outcome of which may be affected by male physiological condition and food availability. Although many mathematical models and empirical studies suggest that the expression of male courtship signals are condition-dependent, decisions about courtship and mating strategies in relation to food availability have not attracted much attention. In this study, we tested whether daily changes in food availability affect males’ decisions about whether to court. We conducted experiments with the fiddler crab Uca lactea by providing males with additional food every other day. In food-supplemented enclosures, males did not increase courtship activity on the days when food was supplemented. However, they built more courtship structures (semidomes) and waved more on the days when they were not given additional food. Male size had a strong influence on the number of days the males courted. We also tested whether the frequency of surface mating, as an alternative reproductive tactic, decreased when food was supplemented. Contrary to our expectation, the number of males that exhibited the surface-mating tactic increased when food was supplemented whereas the number of mate-searching females did not change. Our findings in this field study suggest that reproductive decisions by male fiddler crabs are affected by fluctuating food availability and present body condition, and the alternative mating tactic of this species may be more frequently used by males under good condition.     

Mate availability accelerates male filial cannibalism in a nest brooding fish: effects of number and fecundity of females

Theoretical models predict that increased mate availability accelerates filial cannibalism by the parental male, but we do not yet fully understand how the various aspects of mate availability contribute to this effect. We examined the effects of two elements of mate availability—female fecundity and sex ratio—on filial cannibalism by the lizard goby, Rhinogobius flumineus, which is a paternal nest brooding fish. We used three types of females (stimulus-females): a single female with slim belly (not ready to spawn), a single gravid female (ready to spawn), and two gravid females. Stimulus-females were housed in a transparent cage and shown to subject males just before and after spawning with a separate female, after which males cared for the brood. A single gravid stimulus-female accelerated filial cannibalism, compared with a control-stimulus consisting of an empty cage, but only during the early care period. In contrast, a single slim-bellied stimulus-female did not accelerate cannibalism. A stimulus of two gravid females accelerated cannibalism to the same degree as a single gravid female stimulus. Our results suggest that in lizard gobies, filial cannibalism by parental males is accelerated by female quality (fecundity) in the early care period, but not by a higher number of available females (sex ratio).     

Predation risk affects courtship and attractiveness of competing threespine stickleback males

The effect of predation risk and male-male competition on male courtship behaviour and attractiveness to females was studied in the threespine stickleback (Gasterosteus aculeatus) by presenting dummy or live females to solitary and competing males under different predation risks. In the presence of a predator, males decreased courtship activity. Different courtship components were, however, adjusted to different extents and in opposing directions to predation risk, probably because the single components may have varied in riskiness. The presence of a competing male decreased overall courtship activity, but increased the frequency of zigzags, suggesting zigzagging to be a competitive strategy against other males. In the presence of a predator male courtship activity was not affected by a competitor. Female mate choice correlated with the males' previous frequency of zigzags towards a dummy female. However, when a live female paid attention to a male, the male decreased zigzagging and instead increased leading and fanning behaviours, probably trying to attract the female to the nest to mate. Predation risk affected the attractiveness of males as females reduced their attention to a male when he faced a predator and reduced his courtship activity. As females instead increased their attention to a competing male that had increased his courtship activity, due to decreased competition, males clearly are balancing mating opportunities against predator avoidance. When males vary in their susceptibility to predators, predation risk may thus affect mating success of competing males. Received: 31 January 1997 / Accepted after revision: 15 April 1997     

Courtship and spawning in the emperor angelfish Pomacanthus imperator,with comments on reproduction by other pomacanthid fishes

Courtship and spawning are described for Pomacanthus imperator and 4 Centropyge spp. at Enewetak Atoll, Marshall Islands, observed from 15 August to 15 September, 1981. In all species, courtship and spawning occur at dusk, continue throughout most of the lunar month, are preceded by male display to the female, and culminate in the pair's shedding gametes into the water column. The species differ in ascent height, details of courtship behavior, spawning location relative to prominent reef topography and degree and type of sexual dichromatism. Social organization in all species appears based on male defense of harems of 2 to 5 females. Observed behavior is compared to that of confamilials, and discussed relative to proposed theories concerning lunar periodicity of spawning activity, sexual dichromatism and the adaptive significance of spawning ascent behavior.     

Female choice of multiple male criteria in guppies: interacting effects of dominance,coloration and courtship

Summary I experimentally examined the relative importance of social dominance, color patterns, and courtship behavior in male mating and reproductive success in the guppy Poecilia reticulata. Female choice of males is based on a complex set of behavioral and morphological traits. The results of 59 paired-male one-female visual choice and mating trials showed that male mating success was positively correlated with dominance, courtship intensity, and male coloration. Only dominant males engaged in full copoulations, and they sired two-thirds of the broods. An analysis of the paternity of broods and results of mating trials showed that a female's visual response when the sexes are separated by a glass partition is a good predictor of a male's reproductive success when the partition is removed and they are allowed to mate. A canonical correlation analysis of male behavioral and morphological traits indicated that female visual response and male mating success were positively correlated with male courtship and with agonistic behavior. However, the relative importance of color varied. Carotenoid and iridescent spots were important both in attracting the female's attention and in enhancing male mating success. Melanins were not correlated with either mating success or female response. There was a relatively low correlation (48%) between male behavioral and morphological variables and female response variables (full copulation and female visual response). These results suggest that female choice is subtle, and is based on a complex suite of male behavioral and morphological traits as well as on competitive interactions among males.     

Mating behaviour and reproductive cycle of Archaster typicus (Echinodermata: Asteroidea)

This study examines the reproductive cycle, the mechanism of male-on-female pairing behavior and the spawning behavior of Archaster typicus Müller et Troschel. Field studies were conducted in the intertidal zone of the sand beaches at Penghu, Taiwan (23 32N; 119 33E) at ebb-tide in 1984 and 1985. The pairing behavior of A. typicus is a reproductive behavior which leads to simultaneous spawning, increasing the probability of fertilization. As the breeding season approaches, sea stars, especially males, display increased mobility. Because only males tend to mount another individual and because males can detect the sex of another individual by contact with the side of their arms, a male-on-female pair is formed when a male encounters a female. Eighty-five percent of the sea stars observed were paired during the pairing season. Spawning by a paired female is closely followed by spawning of its paired male; male spawning, however, does not induce spawning in its paired female. During spawning, the male turns slightly so that its arms overlap the arched arms of the female. The gonad volume of males is much less than that of females. This may result from the high efficiency of fertilization in this species, which does not require a large amount of gametes to be released, or from the higher energetic demands made on the males.     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Multiple mating in a lekking bird: why do peahens mate with more than one male and with the same male more than once?

Summary Approximately 50% of marked peahens (Pavo cristatus) mate more than once with lek males. Some females mate with more than one male, others copulate repeatedly with the same male. The frequency of courtship also shows marked variation. Some females repeatedly engage males in courtship interactions after they have succesfully copulated with them. The likelihood of mating with more than one male increases if a female first mates with a non-preferred (unsuccessful male). There is a non-significant tendency for females to copulate with a more successful male when remating. Peahens may mate with a non-preferred male first if they do not encounter a successful male during their initial period of choice, perhaps because the most successful male on a lek was courting another female and/or was defended by another female. There are more aggressive interactions between females in front of preferred males. Preferred males receive more repetitive courtship behaviour and repeated matings. Dominant females are more likely to engage in repetitive courtship and matings. The number of times a female initiates courtship on any one day increases with the number of other females actively courting males at a lek site on that day. We suggest that there is competition amongst females for access to preferred males and that dominant females try to monopolise these males by repeatedly engaging them in courtship interactions. We discuss the implications of these observations for the idea that female may gain directly from mate choice in a species where males contribute nothing but gametes to their offspring. Correspondence to: M. Petrie at the present address     

Strategic male signalling effort in a desert-dwelling fish

Males often use elaborate courtship displays to attract females for mating. Much attention, in this regard, has been focused on trying to understand the causes and consequences of signal variation among males. Far less, by contrast, is known about within-individual variation in signal expression and, in particular, the extent to which males may be able to strategically adjust their signalling output to try to maximise their reproductive returns. Here, we experimentally investigated male courtship effort in a fish, the Australian desert goby, Chlamydogobius eremius. When offered a simultaneous choice between a large and a small female, male gobies spent significantly more time associating with, and courting, the former, probably because larger females are also more fecund. Male signalling patterns were also investigated under a sequential choice scenario, with females presented one at a time. When first offered a female, male courtship was not affected by female size. However, males adjusted their courtship effort towards a second female depending on the size of the female encountered previously. In particular, males that were first offered a large female significantly reduced their courtship effort when presented with a subsequent, smaller, female. Our findings suggest that males may be able to respond adaptively to differences in female quality, and strategically adjust their signalling effort accordingly.     

Prostaglandin F2α facilitates female mating behavior based on male performance

Hormones play an important role in the regulation of reproductive behavior. Here, we examined the effects of the fatty acid derivative prostaglandin F2α (PGF2) on female sexual behavior as well as the interaction between PGF2-induced mating behavior with male courtship display in the lek-breeding African cichlid fish, Astatotilapia burtoni. In a two-way choice paradigm, we found that nonreproductive females preferred to associate with smaller, less aggressive males over larger, more aggressive males. However, PGF2-treated females dramatically reversed their preference to larger males. In a second experiment, PGF2 treatment dramatically increased sexual behavior in nonreproductive females as measured by time spent in the bower of the stimulus male, even when the female and the stimulus male were separated by a transparent divider. This effect was even more pronounced when the stimulus males were exposed to the putative female pheromone 17α,20β-progesterone (17α,20β-P). Under full-contact conditions, only PGF2-treated females visited a stimulus male’s bower, where they even displayed circling behavior usually only seen during spawning. Interestingly, male performance prior to PGF2 treatment predicted female sexual response. Our study demonstrates the importance of PGF2 in the control of female reproductive behavior in interaction with male performance.     

Mating behavior of female rock shrimp Rhynchocinetes typus (Decapoda: Caridea)—indication for convenience polyandry and cryptic female choice

While studies on a wide diversity of organisms have demonstrated the importance of female behavior during matings, in crustacean studies, a strong bias towards male mating behavior prevails. Reproductively mature rock shrimp (Rhynchocinetes typus) exist as several ontogenetic stages that differ in their morphological and physiological capacities. In natural populations, the majority of males are in early ontogenetic stages (termed typus), many are in intermediate stages (intermedius), and few are in the terminal molt stage (robustus). Dominant robustus males, which have already demonstrated their biological fitness by surviving to this stage, have previously been shown to have a higher potential than subordinate typus males to defend receptive females against other males, and fertilize the entire clutch of a female. While females should thus show a preference for robustus males, they nevertheless frequently receive sperm from typus males. These observations suggested that females might have mechanisms to discriminate against sperm from subordinate males. In laboratory experiments, we observed that females avoided being seized by typus males for longer time periods in the absence of robustus males than in their presence. Following seizure, females that were initially held by typus males, required more time to initiate spawning than those held by robustus males. Many typus males transferred spermatophores to females before these started to spawn while robustus males waited until females began to spawn before they transferred spermatophores. Females manipulated spermatophores received from typus males for long time periods (minutes), but not those they received from robustus males. By accepting sperm from subordinate typus males, females may avoid further harassment (convenience polyandry), but they subsequently may discriminate against these subordinate males by delaying spawning and removing their sperm. These observations suggest that female behavior influences the outcome of matings, favoring fertilization of eggs by sperm from dominant males. Convenience polyandry and cryptic female choice may be common in other crustaceans as well.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by T. Czeschlik     

Experience with chemotactile cues indicating female feeding history impacts male courtship investment in the wolf spider <Emphasis Type="BoldItalic">Schizocosa ocreata</Emphasis>

Male fitness is often determined by the ability of the male to gain access to multiple mates, although in species that exhibit sexual cannibalism, males might increase their likelihood of being cannibalized with each encounter. This risk should create selection for males who are able to perceive potential risks associated with mating encounters. We studied male Schizocosa ocreata wolf spiders to determine whether they use female chemotactile cues (silk, excreta) as indicators of potential risks and how these cues affected subsequent male courtship behaviors. Female treatments included satiated females vs. starved females, as well as a treatment where females had recently cannibalized a male S. ocreata. We performed experiments to assess (1) if males use female chemotactile cues to determine potential risks associated with differing female feeding treatments and alter courtship investment, and 2) how male experience with female chemotactile cues affected courtship investment in subsequent female encounters. At first encounter, males do not vary courtship investment (number of bouts, duration, and vigor) with different female feeding treatments. However, male behaviors during subsequent encounters with female chemotactile cues varied in complex ways, depending on female feeding treatment and male first encounter experience. These data suggest that male experience impacts perception of female chemical cues and offers the male opportunity to avoid sexual cannibalism in high-risk situations.     

Sexual selection in the northern pintail (Anas acuta): the importance of female choice versus male-male competition in the evolution of sexually-selected traits

We experimentally studied the relative importance of plumage, dominance status, and courtship behavior in determining male pairing success in the northern pintail Anas acuta and assessed whether these traits function in female choice, male-male competition or both. In an experiment (experiment IA) that eliminated the confounding effects of male-male competition and social courtship, females chose males with pure white breasts and colorful scapular feathers. When the same group of birds were free to interact (experiment 1B), male behavior was more important: females chose males that courted them intensely and were attentive to them, although preferred males again had whiter breasts and more colorful scapulars. In a second experiment (experiment 2), testing the effect of age on pairing success, females showed a significant preference for 2-year-old males over yearlings: 2-year-old males courted more and were more attentive to the female than yearlings; they were also more colorful than yearlings in a number of plumage measurements. Although males (in both experiments 1B and 2) were aggressive to one another while courting the female and dominant males were sometimes able to exclude subordinates from social courtship, contrary to expectation, we found no relationship between initial dominance rank and pairing success or dominance rank and age. In addition, dominance was not correlated with any of the morphological traits measured. Once chosen, however, subordinate males typically initiated fights with the higher-ranked male(s) and quickly achieved dominance. These results suggest that (1) females choose males based on a suite of morphological and behavioral characteristics, (2) male dominance relationships do not constrain active female choice, (3) a male's position in a dominance hierarchy is largely a result rather than a cause of female choice, and (4) female choice plays a more significant role than male-male competition in the evolution of several secondary sexual traits in male northern pintails.     

Influence of the courtship song of the acridid grasshopper Gomphocerus rufus L. on the female

Summary Males of the acridid species Gomphocerus rufus L. perform unusually long courtship. Following their imaginal molt, the behavior of the females changes in three stages: initial rejection of the males, passive acceptance without singing, and active attraction of a mate by singing. Females in the active state permit copulation after brief courtship or even without it.To understand the function of long courtship, two hypotheses are tested by the present experiments. Hypothesis A assumes that the long courtship of the male affects the female so as to accelerate entry into the active mating state. This is not the case: Females raised in isolation enter the active state at the same time as those that have been exposed to courtship continually. Hypothesis B assumes that the long courtship changes the behavior of a female still in the passive state to enable earlier copulation. This hypothesis is tested by longterm recording of the locomotor and singing activity of the female in the presence of a courting male. The records show that the locomotor activity of a female in the passively acceptant state is significantly reduced during male courtship. Moreover, males in the presence of a passively acceptant female court longer than when the female is in either of the two other states.It appears, therefore, that the unusually long courtship serves to keep the female near the male for a longer period of time, increasing the chance of copulation. The advantages and disadvantages of this courtship strategy are compared with those of searching for a female in the active state.     

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