Simultaneous spawning by female stream goby Rhinogobius sp. and the association with brood cannibalism by nesting males

A laboratory experiment was conducted by varying the undersurface area of nesting substratum and the number of females in an experimental tank to elucidate the determinants of the mating pattern in the stream goby, Rhinogobius sp. cross‐band type. Males with larger nests tended to attract two or more females to their nest in a tank. Moreover, males spawned simultaneously with multiple females and entire brood cannibalism by males was rarely observed under a female‐biased sex ratio. When males spawned with a single female with low fecundity, however, entire brood cannibalism occurred at a high frequency, suggesting that a male guarding a nest with fewer eggs consumes the brood. Therefore, spawning behaviour of females that leads to a large egg mass would decrease the risk of entire brood cannibalism. In this species, simultaneous spawning by multiple females in a nest serves as a female counter‐measure against entire brood cannibalism. These results suggest that a conflict of interest between the sexes through brood cannibalism is a major determinant of simultaneous spawning.     

Sound Production by Adult Haddock, Melanogrammus Aeglefinus, in Isolation, Pairs and Trios

Haddock, Melanogrammus aeglefinus, have been previously shown to produce sounds during mating. Several behavioural aspects of sound production of courting haddock were further investigated in relation to sex ratio. We assessed whether (i) single males or females generate sounds when isolated, (ii) sound is produced when one male is present with a female, (iii) sound production becomes altered with the introduction of an additional male, and (iv) sounds are produced independent of egg release. Data were collected from 30 March to 11 June 1999, during the spawning period using small outdoor tanks. Sounds generated by captive males during spawning were categorized as knocks, hums and an intermediate between these two types. Solitary males and females did not produce sounds. Sounds were produced when one male was present with a single female. The knocking call duration increased when a second male was introduced. Sounds produced by males occurred independent of the day of egg release.     

The possibility of sex and nymph discrimination by the male cockroach,Nauphoeta cinerea

The possibility of sex and nymph discrimination by males was investigated in the cockroach,Nauphoeta cinerea (Olivier). A sexually mature male takes a courting position toward a sexually mature female when he comes into contact with her and recognizes her through antennal contact. In contrast, males often behave aggressively toward each other: they bite at each others; wings and/or legs, chase each other and antennate mutually. The male, however, does not show conspicuous behavior (mating behavior or aggressive behavior) toward nearby nymphs. The male produces audible sounds when he courts a sexually mature but non-receptive female who does not respond to his courtship behavior. We found that the male also stridulates after he repeatedly courts immature teneral females, males and (last-instar) nymphs. After the bodies of teneral insects are sclerotized, the male shows courship and stridulation behavior toward sexually mature but non-receptive females but not toward mature males and nymphs. At this stage the male begins to behave aggressively toward other post-teneral mature males. We think that the variability of the sexually mature male's behavior toward other conspecifics (courtship behavior toward female, aggressive behavior toward male and no conspicuous response toward nymph) results from the male's recognition of adult and nymph.     

Differences in Aggressiveness in the Midas Cichlid Fish (Cichlasoma citrinellum) in Relation to Sex,Reproductive State and the Individual

The Midas cichlid, Cichlasoma citrinellum, is monogamous and biparental. Because of competition for limited spawning sites and intense predation on their young, vigorous defense of their territory is essential. Although both sexes engage in defense, they differ in aggressiveness. The aggressive responses of both sexes were measured by counting the number of bites and bumps each fish directed toward its own mirror image. The size of the fish's genital papilla was also recorded to estimate its reproductive state. Compared with females, males had higher median mirror scores with greater variance. The scores of individual males were also more consistent through time than were those of females. Females close to spawning had the highest mirror scores, whereas male scores were highest early in the reproductive cycle. Selection has apparently favored aggressiveness in both sexes. We argue, however, that differences in aggression are the result of selection acting dissimilarly on the two sexes.     

Reproductive behavior of the halfmoon grouper, <Emphasis Type="Italic">Epinephelus rivulatus</Emphasis>, at Ningaloo Reef,Western Australia

Behaviors associated with spawning by the halfmoon grouper, Epinephelus rivulatus, at Ningaloo Reef, Western Australia, are described from in situ observations made each evening throughout most of a lunar cycle. Spawning occurred after sunset on six consecutive evenings during the full moon period. During this time males were particularly aggressive toward one another and maintained high levels of activity among the gravid females that rested within each of their territories. Spawning occurred when a male swam alongside a responsive female and the pair rose in a tight spiral 1–1.5 m into the water column before releasing gametes and returning to the seabed. Spawning activity was followed by a longer nonspawning period (ca. 20 days), when fewer intraspecific interactions were observed and gamete reserves were replenished. Histological and behavioral evidence suggests that this cycle of spawning and replenishment may occur on a monthly basis. Although individual fish, particularly females, moved into certain areas to spawn, E. rivulatus did not form spawning aggregations as do larger species of grouper.     

Male Isaza (Gymnogobius isaza, Gobiidae) prefer large mates: a counterstrategy against brood parasitism by conspecific females

Like many other gobies, male Isaza (Gymnogobius isaza) which are endemic to Lake Biwa, Japan, exclusively care for broods in nests. This goby may have an optimal range of brood size (i.e., an average clutch size of about 2000–3000 eggs) within which they may produce larger numbers of hatching young because much larger broods may be destroyed by fungal infection before hatching. This optimal brood size hypothesis (Takahashi et al. in J Ethol 22:153–159, 2004) predicts that (1) after spawning, both males and females will refuse additional spawning by other gravid females (second females) to keep brood sizes within optimal ranges, (2) larger fish will repel second females more successfully than will smaller fish, and thus, (3) both sexes prefer larger mates. To examine these predictions, we first observed Isaza’s aggressive behaviors in aquaria and investigated whether fish attacked and repelled second females that were introduced after spawning, and, if so, what were the sizes of fish that did so. Large fish, regardless of sex, aggressively prevented second females from entering the nest, but second females larger than the pairs displaced the pair females forcibly and spawned eggs into their clutches. Mate choice experiments showed that males preferred large females. Although females’ choice of large mates was not confirmed, many results may largely coincide with the predictions of the optimal brood size hypothesis. Thus, Isaza males’ choice of large mates will be advantageous for defending against brood parasitism by conspecific females and for achieving optimal clutch size.     

Male Siamese fighting fish use gill flaring as the first display towards territorial intruders

The Siamese fighting fish (Betta splendens) is well known as an aggressive fish with unique spawning and parental care behavior. During reproduction, male fish construct a bubble nest, court females, protect the brood, and defend the territory through aggressive displays. Aggression in male Siamese fighting fish has long been the subject of investigation; however, the kinematics of aggression during contests have been largely overlooked. Here we investigated how nest-holding, male Siamese fighting fish use two different types of displays, gill flaring and fin spreading, towards intruders during various reproductive phases; before (BB) and after bubble nest building, and after spawning (AS), and hatching (AH). Males were more aggressive towards male than female intruders and the level of aggression changed significantly between reproductive phases. Gill flaring, the more energetically costly display, was the dominant initial display towards male and female intruders in BB, AS, AH phases. However, defending males switched to fin spreading after prolonged exposure to intruders. The results suggest that Siamese fighting fish use gill flaring as an acute response in order to defend their territory; this response may be replaced by fin spreading as a chronic response, probably to reduce the energetic costs during the contest.     

Maternal aggression and intermale social aggression: a behavioral comparison

The aggressive behavior of alpha male rats and lactating females were each examined toward an intact adult male rat, a castrated adult male rat, an anesthetized adult male rat, a nonlactating adult female rat, an adult albino guinea pig (male or female), or an albino mouse (male or female). When in their living colony, females displayed high levels of aggressiveness toward all stimulus objects except a mouse. The aggression toward the intruding males occurred whether the female's pups were present or not. Alpha males were aggressive toward the same stimuli except an intruding female rat and a mouse. When tested in an unfamiliar colony, the males but not the females (with or without pups present) were aggressive toward an adult male rat. Half of the females but none of the males displayed defensive burying toward an anesthetized intruder. It is suggested that the attack on an adult female, the absence of attack outside of the resident colony, and the tendency to display defensive burying are features of the aggressiveness of lactating females that are fundamentally different from the aggressiveness of alpha males. The form of the aggression (lateral attack vs. lunge attack) was only quantitatively different in males and females.     

Male competition and female choice interact to determine mating success in the bluefin killifish

Whether male competition and female choice act in concert, independently,or in opposition is a critical issue for understanding sexualselection. In complex social systems, the outcomes of pairwiseinteractions may not be accurate indicators of how sexual selectionemerges. We investigated how female choice and male competitioninteract in the bluefin killifish, Lucania goodei, in a 3-stagedexperiment where 1) females could choose between 2 males, 2)those males could interact in the presence of that female, and3) females and males could freely interact and spawn. In thepairwise stages (1 and 2), females displayed pronounced preferencesbetween males and male competition produced a distinctly dominantindividual. None of the morphological traits, including color,measured in males were associated with either female preferenceor male dominance. When all 3 fish interacted (stage 3), maleactivity level was the sole predictor of spawning success. Maleswith elevated activity levels were more aggressive toward malesand females, exhibited intensified courtship, and obtained morespawns. Female preference did not predict the number of spawnswith a male, but it did predict her latency to spawn; femalesspawned more quickly with preferred males. Thus, male competitionand female choice interact to determine reproductive success,but there is evidence for conflict and a cost to females ofassociating with dominant males. Reproductive success in thisspecies is not easily predicted from simple measures of morphologyor female preference and is influenced by complex social interactions,both between males, and between males and females.     

Some relationships between the process of pair formation and gamete maturation in Harmothoë imbricata (L) (Annelida: Polychaeta)

Male and female Harmothoë imbricata form pairs in preparation for spawning although the individuals are normally mutually antagonistic. This change in behaviour is due to a reduction in aggression shown by the females as they approach maturity. Immature males attempt to mount but are normally repulsed by the aggressive immature females. No obvious behavioural change occurs during the period of final gamete maturation in the male. Pair formation results in the paired female spawning sooner than would an equivalent female kept in isolation. Release of sperm over the newly formed egg mass is ensured by receptors on the ventral cirri which inform the male that the female has spawned.     

Sexual dimorphism of drumming muscles in European hake (<Emphasis Type="Italic">Merluccius merluccius</Emphasis>)

Dissections of mature and non-mature European hake males and females (N = 142) collected in waters off the western coast of Norway and in the Bay of Biscay (France) in 2004–2006 demonstrate for the first time that this gadoid species contains drumming muscles. There were differences in drumming muscles weight with body length, sex and maturity stage. This study shows that the difference between females and males is primarily manifested during the spawning season, seen both in the French and Norwegian samples. For the mature females the drumming muscles dry weight increases only slightly, if at all, with increase in total length. For mature males there is a corresponding rapid increase. There does not seem to be any consistent difference between the average dry weight of the drumming muscles in immature male and immature and mature female hake of the same length, tested on the Norwegian samples. Our results suggest that male hake, like the males of other gadoids studied, may produce sounds in the context of spawning.     

Sound production and reproductive behaviour of the armoured catfish Corydoras paleatus (Callichthyidae)

Sound production during reproductive behaviour, dyadic encounters and distress situations was investigated in the callichthyid catfish Corydoras paleatus. Sounds were broad-band, pulsed, acoustic signals produced during abduction of the pectoral spines. Only males emitted trains of sounds during courting and trains of sounds of shorter duration during dyadic encounters. Several males, which are usually smaller than females, courted one gravid female without obvious cooperation or competition between them. During mating, one previously vocalizing male clasped the female's barbels with one pectoral spine and inseminated the eggs. The number of successful spawnings, days until spawning, and number of eggs laid was not related to the number of males (one, two or three) combined with one female. Males did not behave aggressively towards each other during courting or in dyadic encounters. In distress situations, when fish were hand held, both sexes and juveniles produced single sounds. The dominant frequency was negatively correlated with body size and the sound duration was positively correlated with relative length of pectoral spines (standardized to body length). This acoustical behaviour in C. paleatus differs considerably from Hoplosternum thoracatum, a representative of the callichthyine subfamily, in which vocalization was observed during territorial behaviour in males and aggressive behaviour in both sexes. This is the first report of a major difference in vocalizing behaviour within one teleost family.     

Aggressive Behavior and Courtship Rejection in Brightly and Plainly Colored Female Keeled Earless Lizards (Holbrookia propinqua)

Relationships between bright secondary sexual coloration and behavior were studied in female Holbrookia propinqua, which develop striking orange and yellow colors during the breeding season. In tethered introduction studies, brightly colored females performed aggressive courtship rejection behaviors toward conspecific males; plainly colored females were not aggressive toward males, but attempted to avoid them. Responses of females of the two color patterns toward conspecific females of both color phases were not detectably different. Experimental introductions of lizards with coloration modified by paint showed that females of both color patterns recognize any other lizard bearing the bright female colors as female, regardless of actual sex. Both the orange and yellow components were shown to contribute to sex recognition. The yellow component alone allowed accurate sex identification, but only half the females responded to males painted with only the orange female component as if they were females. Because females did not behave differently toward other females on the basis of coloration, the hypothesis that bright coloration evolved as an adaptive signal between females is rejected. The dark ventrolateral stripes of male and plainly colored females did not appear to affect intraspecific social responses by females.     

Changes in behaviour associated with pair formation in the polychaete Harmothoë imbricata (L.)

Spacing between individuals in populations of Harmothoë imbricata has been investigated both on the shore and in the laboratory. Males tend to occur closer together than females, and the mean male‐male individual distance measured was less than the mean distance between females; male—female distances for immature worms were intermediate. When worms mature they pair: the male mounts the female and lies across her dorsal surface. There is evidence that after spawning the members of a pair separate. Contact responses between worms have been investigated in the laboratory. Most encounters between immature worms lead to separation, as a result of one or both worms moving rapidly away from the other or fighting; females show more marked avoidance behaviour than males. The majority of male‐male and female‐female encounters between mature worms also lead to separation but in male‐female encounters the male usually mounts the female. A male which has mounted a female becomes highly aggressive and will attack intruding males but not females.     

Relatively large males lower reproductive success in female zebrafish

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

Reproductive strategy and interspecific competition in the Lake-living gobiid fish isaza,Chaenogobius isaza

Isaza,Chaenogobius isaza, is a small gobiid fish endemic to Lake Biwa. It lives offshore throughout almost the entire year, showing a remarkable diel vertical migration. In early spring, males and females migrate to lake shore to spawn under stones. Spawning season is limited to a very short span of time in early spring, late April to early May. During this short spawning season, the male is supposed to have only 2 brood cycles at maximum. At each brood cycle, the male is strictly monogynous, never accepting additional females. Males therefore show a marked mate choice, choosing a larger female regardless of the size of the male himself. Females also choose larger males. However, males are supposed to not waste time in male-male fighting in the natural spawning area. That this very short spawning season and its occurrence in early spring is primarily to avoid interspecific competition with another littorally reproducing goby Yoshinobori,Rhinogobius brunneus, is experimentally demonstrated.     

Optimal Mate Searching Strategy in the Symbiotic Spider Crab Inachus phalangium (Decapoda)

The reproductive strategy of the symbiotic spider crab, Inachus phalangium was studied from 1981 to 1983 in the natural environment. Main study focus was the efficiency of males in search of mates. I. phalangium females are site-constant, and live in the protection of one anemone or group. Males travel frequently between anemones harbouring females due to spawn; they copulate and guard the females until spawning, after which the male leaves again. A male operates in a patrol area containing 3–8 anemone groups and up to 8 females, visiting each female in turn repeatedly just before it is due to spawn. Patrol areas of different males may overlap, with resulting competition to fertilize a female's next brood. Large males have higher reproductive success than small ones. Females live up to 8 months after the moult of puberty and hatch up to six broods, and males live up to 7 months as adults. A male could fertilize a calculated 26,000 eggs, whilst a female's reproductive potential is ca. 4,200 eggs. Mortality risks are higher for males than for females, probably because of increased predation while leaving the protection of anemones in order to visit females. Males learn the positions of anemones harbouring females in their patrol areas, and when these are due to spawn. This allows a male to travel with a target and arrive punctually to fertilize the next brood due in his circuit. I. phalangium is the first marine invertebrate reported to use a “schedule” of localities and times for visiting prespawning females. In this way males minimize searching time and mortality risk, and maximize the number of broods fertilized.     

Mate Choice and Spawning Success in the Fighting Fish Betta splendens: the Importance of Body Size, Display Behavior and Nest Size

Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer‐generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non‐displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer‐generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer‐generated females that performed courtship displays over non‐displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size‐assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species‐recognition function.