The nest-holding grass goby (Zosterisessor ophiocephalus) male adjusts the spawning activity in relation to parasitic nest intrusions

This study examines the temporal pattern of spawning behavior by the territorial (i.e. nest-holding) grass goby male, Zosterisessor ophiocephalus, in response to sneak intrusions by the small parasitic male under controlled laboratory conditions. The spawning activity of the territorial male consists of a sequence of upside–down movements on the ceiling of the nest accompanied by undulations of the body and sperm release. Five pairs of one territorial male and one parasitic male, each kept inside a large tank provided with an artificial buried nest (always occupied by the territorial male) and one small tunnel-shaped shelter (always occupied by the small male), were observed during one-female spawning taking place in the innermost part of the nest (i.e. the nest chamber). During the spawning, the presence of the small male nearby the nest openings elicited aggressive behavior and increased nest patrolling by the territorial male. In one spawn the small male never attempted to enter the nest. In four spawns the small male entered one to three times the nest chamber (sneaks), staying there from 2 to 203 s until the large male chased him away. The temporal patterning of the spawning activity by the territorial male (bouts of upside–down, U–D), and its changes following a sneak by the small male, were investigated using bout analysis and correlative tests. Results showed the length of bouts U–D did not change significantly after a sneak intrusion. whereas gap length (i.e. the period between subsequent bouts U–D) decreased significantly after each sneak intrusion. The mean duty cycle of bouts U–D tended to be positively correlated to the number of sneaker intrusions of each replicate. Results are discussed in the light of current knowledge on sperm competition among externally fertilizing teleosts.     

Electric signalling and reproductive behaviour in a mormyrid fish, the bulldog Marcusenius macrolepidotus (South African form)

Bulldog fish (Marcusenius macrolepidotus) generate short (<1 ms) electric-organ discharges (EODs), separated by much longer and highly variable interdischarge intervals (IDIs). We observed overt behaviour and electrical activity during reproductive behaviour in a male and in a female bulldog, and identified IDI patterns with putative signal functions. In contrast to Pollimyrus adspersus and Pollimyrus isidori, in which an elaborate and extended courtship precedes spawning proper, our fish started spawning almost immediately when we allowed the female to enter the males territory. The male did not construct a nest, and neither parent provided parental care. The male showed very little aggression towards the intruding female. Fish spawned in bouts near the males hiding place, and eggs were scattered by the females vigorous tail flips as she left the spawning site, only to return shortly thereafter. During spawning bouts, both fish generated highly stereotyped IDI patterns: the male generated a series of IDIs gradually decreasing from about 200 ms to about 55 ms that was abruptly terminated by a long IDI. The female generated a series of relatively regular IDIs (about 54 ms) that was followed by a marked increase in IDI duration (the probable time of spawning). Finally, a sharp decrease in IDIs to about 20 ms accompanied the females sudden escape from the spawning site. In between spawning bouts, both fish generated series of very short IDIs (high discharge rate, HD) that alternated abruptly with very low-rate inter-HD activity (especially in the male). IDIs as short as 9 ms (male) or 11 ms (female) occurred during HD displays. No visible aggression, in fact very little overt behaviour, occurred during these HD displays in both fish. Agonistic interactions between male and female, outside a reproductive context, were similar to those previously described in male pairs, including overt behavioural patterns such as parallel swimming, antiparallel display and attack, as well as HD displays. When not interacting, fish did not generate HD displays. We suggest the HD display is a communication signal in both reproductive and agonistic contexts.     

Predation on eggs by conspecific males in two Japanese damselfishes,Pomacentrus nagasakiensis andChromis notatus notatus,after removal of egg-guarding males

Egg predation by conspecific males of damselfishes,Pomacentrus nagasakiensis andChromis notatus notatus, was studied at Bohnotsu (31°15′ N, 131°15′ E) and Tsuyazaki (33°47′ N, 130°29′ E), respectively. InP. nagasakiensis, 7 egg-guarding males were removed from nests, and intrusion and egg predation by conspecific males took place in all of the nests within an average time of 12.2 min. Frequency of attacks on eggs by other reef fishes during the period between the removal of egg-guarding males and intrusion of conspecific males was lower than expected, considering the frequency of aggressive displays of the egg-guarding males. The results of male removal experiments forChromis notatus notatus were almost the same, with intrusion and egg predation by conspecific males occurring in all 12 of the nests from which egg-guarding males had been removed. SixP. nagasakiensis and 15C. notatus notatus which intruded into the nests and preyed on eggs were collected. All were males except for 1C. notatus notatus female. Results suggest that these males which intruded into vacant nests to eat eggs did not have clutches of their own during the spawning bouts when the experiments were done. Results also showed that the intruding males were not inferior to egg-guarding males in standard length, gonadosomatic index, and condition factor. It is clear that males which do not have their own clutches during at least 1 spawning bout and prey on others' eggs when egg-guarding males disappear are common in these 2 gregariously nesting damselfishes. It is discussed that conspecific eggs are potentially a good food source not only for other species of reef fishes but also for conspecific males which do not have their own eggs during the spawning bouts. The origin of males wich prey on eggs of other conspecifics is also considered. Some aspects of egg-guarding and aggressive behavior of males in damselfishes are discussed.     

Pelagic spawning of the hawkfish Oxycirrhites typus (Cirrhitidae)

Synopsis Pelagic spawning of the deep slope coral-dwelling cirrhitid Oxycirrhites typus was observed for two social groups at Papua New Guinea. This species was previously reported to be a demersal spawner in an aquarium. Courtship in social groups consisting of a single male and one or two females commenced just prior to or after sunset among the branches of gorgonian or antipatharian corals. Males and females occupied separate corals; males either visited females at their corals or met them at an adjacent coral just prior to courtship. Courtship was sequential and consisted of two or more bouts with each female that culminated in a rapid ascent into the water column and the release of floating eggs. Fertilized eggs, taken from a third social group, were spherical and averaged 0.69 mm in diameter. Spawning pairs sought refuge in their resident corals or in the coral where courtship occurred immediately after spawning was completed.     

Female Leadership During Migration and the Potential for Sex-Specific Benefits of Mass Spawning in the Brown Surgeonfish (Acanthurus nigrofuscus)

Spawning aggregations in coral-reef fishes have been hypothesized to confer any one of several mutually non-exclusive benefits, largely expected to serve the interests of both sexes simultaneously. Here we provide indication that in the brown surgeonfish, Acanthurus nigrofuscus, spawning aggregations may confer a sex-specific benefit. Following tagged individuals en-route to their daily spawning-aggregation site we found that while migrating groups (≤20 fish) consist of both males and females, females tend to occupy the lead position more often than expected by chance. In addition, we found evidence that female A. nigrofuscus divide their daily egg-stock among several spawning bouts within the aggregations. We propose that female leadership en-route to spawning aggregations, together with the potential benefits of multiple female mating, are consistent with a sex-specific benefit to spawning aggregations.     

Reproductive biology of racer goby Neogobius gymnotrachelus in the Włocławski Reservoir (Vistula River, Poland)

Initial studies on the reproductive parameters and spawning behaviour of racer goby Neogobius gymnotrachelus, one of three Ponto‐Caspian gobiid species recently invading Polish waters, were carried out in W?oc?awski Reservoir (lower River Vistula, Poland), where spawning lasted from April to August. Males were on average about 23.3% longer and 53.8% heavier than females. The smallest female with mature gonads was 58 mm TL (estimated age 2+ years), and males were sexually mature at 59 mm TL (age of 1+ years). Gonado‐somatic index at the beginning of spawning was seven times larger in females (mean = 8.4) than that in males (mean = 1.2). Mean absolute fecundity was 952 eggs per female (361–2236). Egg diameter ranged from 0.140 to 1.558 mm, being of bi‐modal distribution, which suggests two or three spawning bouts. Length–frequency analysis of juveniles revealed a multimodal distribution. Specialized reproductive behaviour (nest construction and parental care of eggs) and an extended spawning period increase the probability that racer goby can successfully establish self‐sustaining populations in novel environments.     

Sperm traits in relation to male quality in colonial spawning bluegill

Sperm traits (morphology, motility and concentration within ejaculates) and various correlates of male quality (age, body condition, spawning location and timing) were studied in bluegill Lepomis macrochirus , breeding in both the interior and periphery of six colonies in Lake Opinicon, Ontario, Canada. Sperm traits varied significantly more among than within males suggesting that some aspect of male phenotype might influence sperm morphology and behaviour. No measures of male body condition or size were correlated with any sperm or ejaculate traits, when controlling statistically for confounding variables. Sperm swimming speed increased significantly with male age and varied significantly among spawning bouts (controlling for sperm tail length) suggesting that some unknown aspects of male quality might influence the fertilization capacity of spermatozoa. Sperm concentration in ejaculates was significantly higher in males nesting in the interior rather than the periphery of a colony suggesting that those males might also have higher fertilization capacity correlated with their superior dominance status or the lower risk of sperm competition. Thus, older males nesting in the interior of a colony during the first spawning bout of the season are expected to be the best sperm competitors in this population, but the physiological reasons for this increased fertilization capacity remain unknown.     

Reproductive life history of the introduced peacock grouper Cephalopholis argus in Hawaii

This research investigated the reproductive biology (sex ratio, hermaphroditic pattern, size and age at maturity) of Cephalopholis argus, known locally in Hawaii by its Tahitian name roi. The results suggest that C. argus exhibits monandric protogyny (female gonad differentiation with female to male sex change) with females reaching sexual maturity at 1.2 years (95% c.i .: 0.6, 1.6) and 20.0 cm total length (L_T; 95% c.i .: 19.6, 21.2). The female to male sex ratio was 3.9:1. The average age and L_T at sex change was 11.5 years (95% c.i .: 11.1, 12.9) and 39.9 cm (95% c.i .: 39.5, 41.2), respectively. Current information on spawning seasonality of this species is incomplete, but based on the occurrence of spawning capable and actively spawning females, spawning probably takes place from May to October. Evidence of lunar spawning periodicity was found, with an increased proportion of spawning capable and actively spawning females, and an increased female gonado‐somatic index during first quarter and full‐moon phases. This information fills a valuable information gap in Hawaii and across the species' native range.     

Reproductive Success Associated with Territoriality, Sneaking, and Grouping in Male Rose Bitterlings, Rhodeus ocellatus (Pisces: Cyprinidae)

The spawning success of male rose bitterlings, Rhodeus ocellatus, adopting an alternative reproductive style, was estimated through behavioural data and electrophoretic paternal analyses in field observations and experiments. Three mating patterns were observed: territoriality, sneaking, and grouping. Mating patterns depended on a male's relative size and on local male density (the number of males around a spawning spot: a mussel). Spawning patterns (pair spawning, pair spawning with sneaker, and group spawning) varied with local male density. Time-budget data of the territorial males indicated a trade-off between chasing and courtship behaviour as local male density changed. Females deposited appoximately only 1 egg per egg-laying into the mussels. As a result of isozyme analysis, a minimum of 12% (two out of 17) of the offspring in the sample were found to have been fathered by sneaker males in pair spawning with sneaker. I scored through behavioural data the mating success per spawning for each pattern, on an individual basis. The average reproductive success per spawning for each pattern was: territorial (0.61), sneaking (0.31) and grouping (0.11), and thus the successes of the patterns were not equal. Accordingly, the alternative reproductive styles of male rose bitterlings are best interpreted as alternative phenotypes in a conditional behaviour.     

Sperm release behaviour and fertilization in the grass goby

Nesting males of the grass goby Zosterisessor ophiocephalus showed bouts, with intervals of c. 30 min duration, of upside-down movements while rubbing its genital papilla onto the ceiling of its burrow. Such behaviour was shown during female courting and spawning, and even after female removal. Observations showed that this behaviour was associated with the release of a sperm trail on the substratum and clumped spermatozoa in water, agglutinated with a mercury-bromophenol blue and PAS positive glycoprotein secretion of the sperm duct glands. Agglutination in the secretion delayed the activation of spermatozoa and contributed a steady supply, for up to 40 min, of motile spermatozoa during prolonged egg laying by the female. Sperm released before egg laying achieved c. 50% fertilization success compared with nearly 100% obtained if the sperm was released during egg laying. The sperm release behaviour may improve the nest owner's reproductive success against intruders or sneakers. It also allows defence of the nest while the female is spawning and may allow the male to court other females in the proximity.     

Large and persistent effect of a female steroid pheromone on ejaculate size in goldfish Carassius auratus

This study determined if ejaculate size in male goldfish Carassius auratus is increased by the female preovulatory steroid pheromone 4‐pregnen‐17,20β‐diol‐3‐one (17,20βP), which previously has been shown to affect male behaviour and to increase sperm motility and stripped sperm number, and also to increase paternity in competitive spawning and competitive in vitro fertilization. Experimental males were exposed overnight to 17,20βP whereas control males were not. The morning following exposure, each male was placed with a reproductively active female and, after one to 20 spawning acts, aquarium water was sampled to quantify released sperm. Although exposure to 17,20βP induced a five‐fold difference in the number of sperm that could be stripped, the median number of sperm in first ejaculates of pheromone‐exposed males was >60 sixty times that of control males, a pheromonal effect on ejaculate size that persisted for at least 20 spawning acts. The magnitude of the pheromone effect on ejaculate size indicates that it is a critical component of C. auratus sperm allocation, and that examining this effect in concert with other factors (e.g. presence of competitors, male and female size and frequency of spawning) will reveal the contribution of the preovulatory pheromone to male fitness in this promiscuous species.     

The Sexual Behaviour of the Great Crested Newt,Triturus cristatus (Amphibia: Salamandridae)

The sexual behaviour of Triturus cristatus is described. The first stage of sexual behaviour consists of an orientation phase in which the male approaches the female, sniffs her and moves in front of and perpendicular to her. The second stage is a prolonged period of static display with two different tail movements, called Fan and Lash, in 13.8 ± 1.3 bouts (x? ± SE) with intervals of 10.5 ± 0.3 s between bouts. Each bout contains 6.7 ± 0.3 regular Fan beats at a frequency of 0.5–0.8 Hz. This Fanning provides olfactory and mechanical stimulation to the female. 15% of bouts also contain a violent Lash, in which the male rapidly slaps the female's flank. Some Lashes are used to stop the female when she tries to move away. In the third stage of the sexual sequence, the male turns away, Creeps for 6.7 ± 0.3 s (x? ± SE) and then Quivers his tail. A receptive female follows and touches his tail, and the male deposits a spermatophore and then turns to one side with his tail folded along his flank in a position called Brake. The female now approaches again and the male steps away sideways in a movement called Rebrake. After several Rebrakes the male stops, the female touches his tail and the spermatophore may now be picked up in her cloaca. This sequence can be repeated several times during one encounter. This behaviour is discussed in relation to previous studies of courtship in this genus.     

Courtship and spawning behavior of the pygmy grouper,Cephalopholis spiloparaea (Serranidae: Epinephelinae), with notes on C. argus and C. urodeta

Synopsis Courtship and spawning behavior is described for the grouper Cephalopholis spiloparaea from Rota, Mariana Islands. Daily courtship behavior began late in the afternoon and proceeded until after sunset. Males repeatedly visited females in single-male, multiple-female mating groups during each period and engaged in courtship bouts. Mating was observed only twice, was paired and pelagic. Two congeners, C. argus and C. urodeta, with overlapping geographical distributions, were also examined. Cephalopholis urodeta appeared to have a mating group organization similar to that of C. spiloparaea, with some similarity in courtship behavior. Cephalopholis argus also had a similar mating group organization but differed in courtship behavior. This species' behavior more closely resembled that of similarity-sized Epinephelus spp.     

Large schools of Japanese sardines,Sardinops melanostictus, mate in single pair units at night

Synopsis We observed the behavior of the spawning group of Japanese sardine,Sardinops melanostictus, in the field at night, and also confirmed the spawning behavior in the darkness in captivity. The chasing and mating were done in a pair of 1 male and 1 female. The 1-to-1 mating was a definite finding irrespective of the group density in the field, the density in the tank in captivity, or the sex ratio. This maybe because there is little visual information during spawning at night.     

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.     

Female mate choice through spawning parade formed by male-male competition in filefishRudarius ercodes

In the small filefishRudarius ercodes inhabiting temperatre seaweed beds, 2–10 males follow a gravid female, in a line until spawning early in the morning. Formation of the spawning parade and change of male order were investigated in a large aquarium. Males competed for the position nearest to the female during the parade, while the female controlled the parade by her swimming speed and course. During the parade, male members and their order changed frequently. The male order in the parade was related with the intensity of nuptial coloration, which changed according to the social rank. The female sometimes tried to take a spawning posture but rejected spawning when male positioning was not good for her, for example, far different from parading order. The female spawned with 2–5 males on each side. The nearest mating positions to the female were usually occupied by two males that had been staying at an anterior position in the parade for a long time. During the spawning parade over a long distance and time, females ofR. ercodes can choose multiple suitable males in a variable environment like a temperate seaweed bed.     

The song structure and seasonal patterns of vocal behavior of male and female bellbirds (Anthornis melanura)

The bellbird (Anthornis melanura) is a honeyeater endemic to New Zealand, which uses song to defend breeding territories and/or food resources year round. Both sexes sing and the song structure and singing behavior have not yet been quantified. The number of song types, spectral structure, repertoire size, and singing behavior of male and female bellbirds was investigated for a large island population. Song types differed between the sexes with males singing a number of structurally distinct song types and females producing song types that overlapped in structure. Singing behavior also differed between the sexes; males often sung long series of songs while females sung each song at relatively long and variable intervals. Singing by both sexes occurred year round but the frequency of male and female singing bouts showed contrasting seasonal patterns. The frequency of female singing bouts increased as the breeding season progressed, whereas male singing bouts decreased. In contrast to almost all studied passerines, female bellbirds exhibited significant singing behavior and sung songs of complex structure and variety that parallel male song. These results provide a quantitative foundation for further research of song in bellbirds and in particular the function of female vocal behavior.     

Spawning behavior of Arctic charr (Salvelinus alpinus): Spawning synchrony,vibrational communication,and mate guarding

A mismatch in synchrony between male and female gamete release in external fertilizers can result in reduced or failed fertilization, sperm competition, and reduced paternity. In Arctic charr (Salvelinus alpinus), males can adopt either a guard or sneak tactic resulting in both pre‐ and postcopulatory competition between males with alternative reproduction tactics. Here, spawning behavior of free‐living Arctic charr was video‐recorded, and their reproductive behavior was analyzed. From evaluating 157 spawning events, we observed that females mainly spawned with a guarding male and that the female and the guarding male synchronized timing of gamete release under sperm competition. Although sneakers spawned with higher synchrony than the guarding male in single‐male spawning events, the average sneaker released his milt less synchronized with the female than the guarding male under sperm competition. Approximately 50% of the recorded spawning events occurred under sperm competition, where each event included an average of 2.7 males. Additionally, sneakers were more exposed to sperm competition than guarding males. An influx of males, in close proximity to the female, occurred during the behavioral sequences leading up to egg release, but this influx seemed not dependent on egg release, suggesting that something else than gonadal product attracts sneaker males to the spawning female. Just before and during the actual release of gametes, the spawning couple vibrates their bodies in close contact and it seems likely that this vibrational communication between the spawning couple, which results in a larger amplitude sound wave than seen under regular courting, reveals time of gamete release to sneaker males. Thus, vibrational communication may enable synchrony between the guarding male and the female, and this might be traded against the cost of higher detectability from surrounding sneaker males, eavesdropping in close proximity.     

Spawning behavior ofChaetodon multicinctus (Chaetodontidae); pairs and intruders

Synopsis Spawning by the banded butterflyfish,Chaetodon multicinctus (Chaetodontidae) was observed on coral reefs off Kona, Hawaii. These fish occurred in male-female pairs during normal daytime activities, a behavior which is typical for the family. Courtship is also a paired male-female activity. During spawning, however, other individuals (males?) may intrude on the spawning pair. Spawning typically takes place at least a meter or two above the bottom. The spawning position consists of the male below and behind the female with his snout against the female's ventral flank or anal fin area. Intruding individuals may join in when the pair is in position and about to spawn. Intruders line-up against the male in the same position as he is against his female. Underwater photographs are included to illustrate these behaviors.     

Mating systems of two midwater-spawning cichlids,Cyprichromis microlepidotus andParacyprichromis brieni,in Lake Tanganyika

The breeding habits of 2 maternal mouthbrooding cichlids,Cyprichromis microlepidotus andParacyprichromis brieni, were investigated in Lake Tanganyika. Although spawning on the substrate in the male's mating territory is prevalent in maternal mouthbrooders, bothC. microlepidotus andP. brieni spawned in the water column. MaleC. microlepidotus established their mating territories in the open water column, while maleP. brieni did so around fixed spawning sites near a vertical rock surface. In both species, females visited male mating territories, departing soon after spawning and collecting the eggs. Sneaking, which was observed only inP. brieni, may be attributed to the presence of refuges for sneakers in this species. FemaleC. microlepidotus deposited their entire clutch of about 9 eggs in one male territory. In contrast, femaleP. brieni divided their clutch of about 11 eggs among several males. After the final egg-release, femaleC. microlepidotus repeatedly approached their mate, with the mouth near the abdomen of the latter (nuzzling), but femaleP. brieni often departed without nuzzling. Males may eject sperm during nuzzling to fertilize eggs inside the female's mouth. However, maleP. brieni is also known to eject sperm near spawning females when the females are not nuzzling. Such behavior seems to be a male countermeasure against female mate infidelity, because males could not ensure paternity of eggs by ejecting sperm only during female nuzzling.