Reproductive behaviour of the filefishRudarius ercodes: male spawning parades and female choice

Synopsis The reproductive behaviour ofRudarius ercodes is described from undersea observations in Aburatsubo Bay, Japan. Reproductive behaviour can be separated into four parts: (1) Prespawning Search = searching for spawning sites by females and searching for gravid females by males, (2) Spawning Parade = males follow a gravid female in a line, competing with each other to reach the head of the queue, (3) Spawning = the female takes the spawning position, males rush to the side of the female, and mating occurs between one female and several anterior males of the spawning parade, (4) Parental Care = females attach adhesive eggs to seaweed with the mouth and guard them until embryos hatch. There is no male parental care. The reproductive season ranges from May to October and spawning occurs early in the morning every day. Females begin feeding early in the morning, but males feed little at this time.R. ercodes shows neither territorial behaviour nor fixed-pair spawning. One male might spawn several times in one morning. One female spawns at most once every 5 days. The mating system of this species is promiscuous. The probable function of the spawning parade as a style allowing female choice is discussed.     

Spawning behavior of the kissing loach (Leptobotia curta) in temporary waters

The natural spawning behavior of the kissing loach, an endangered species of Botiidae, was investigated in the wild in early June for two years in relation to several environmental factors. Kissing loaches spawned in temporary waters after elevation in water level. All spawnings observed (n=163) occurred within 3-5.5 hours from late afternoon to night after formation of the temporary water. These spawnings were performed by one female and one (71%) or two (29%) males in densely vegetated lentic waters. The female and following male(s) swam into dense grasses, where they vibrated to spawn intermittently. After the vibration continuing for 3-20 seconds, they moved to other parts of the dense grassy area and began vibration again. This sequence of spawning behavior was usually repeated several times, and the eggs were thus scattered widely. The spawning behavior and the rapid larval development of this species appear to be adaptations for the use of temporary waters as a spawning ground. The rise in water level and the consequent formation of temporary waters appear to be crucial triggers for reproduction of the kissing loach.     

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.     

The spawning behaviour of the endangered Eastern Cape rocky, Sandelia bainsii (Anabantidae), in South Africa

Sandelia bainsii is an endangered freshwater fish species which only occurs in short sections of several Eastern Cape river systems in South Africa. The spawning behaviour of S. bainsii is described for the first time. Female S. bainsii remained a light grey colour. Males exhibited distinctive nuptial coloration consisting of an intense black with a white bar on their caudal fin and white windows on the posterior areas of their dorsal and anal fins. There was a darkened post orbital contact organ of ctenoid scales on the reproductively active males. The spawning behaviour was recorded on a video recorder and analyzed during frame by frame replay. Sandelia bainsii are uniparental substrate breeders. The male prepared a spawning area by clearing the substrate with rapid body movements (sweeps). There were a number of preliminary courtship movements, several trial embraces by the male before the actual final embrace. The male underwent a U bend around the female, anchored himself with the contact organ, squeezed the abdominal area of the female with a quick head movement. Courtship and spawning all took place near the substratum. Twenty one separate mating bouts were filmed and analyzed. The duration varied from 1.44 to 5.11 seconds with an average embrace of 3.39 seconds. During the mating clench the female was not normally turned over, as in some anabantoids, but remained upright. The male then guarded the spawning area. Eggs were light yellow, adhesive, demersal and 1.28 mm in diameter with 0.61 mm oil globules. The breeding behaviour of S. bainsii has some similarities with both breeding behaviourial groups, the bubble nesters and the free spawners, within the other African anabantid genus, Ctenopoma. Sandelia bainsii have acquired some temperate specializations with regard to their reproductive style. Free spawning and the release of buoyant eggs is the plesiomorphic anabantid reproductive style. The demersal eggs and male substratum nest sites of S. bainsii are considered apomorphic.     

Precopulatory mate guarding and mating behaviour in the rotifer Epiphanes senta (Monogononta: Rotifera)

Epiphanes senta is a littoral rotifer species that occurs in temporary waters and displays a mating behaviour which has not, to my knowledge, so far been described for monogonont rotifers. Monogonont rotifers show distinctive periods within their life cycle during which mictic females appear. Mictic females produce haploid eggs that develop into males or into diapausing eggs if fertilized. The females of E. senta are mostly stationary on the substrate while males are more active swimmers. If they encounter eggs with female embryos of their own species, they attend them and mate with the hatching female. Experiments showed that males are able to discriminate between male, female and diapausing eggs. They exhibit a strong preference for female eggs that are only a few hours away from hatching compared with eggs in early developmental stages. Further experiments did not show any significant differences in male attendance of mictic and amictic eggs. It is hypothesized that males judge the age of a female egg by sensing a chemical that is produced by the growing embryo and diffuses through the egg shell. The male mating behaviour is similar to precopulatory mate guarding known from arthropods but it lacks the monopolization of the female by the male.     

Spawning and subsequent copulating behavior of the elkhorn sculpinAlcichthys alcicornis in an aquarium

The spawning and subsequent copulating behavior of the elkhorn sculpin (Alcichthys alcicornis) were observed in an aquarium. Soon after the female was introduced into the tank, the male approached and initiated courtship behavior by unfolding all fins. Females which carried ovulated eggs eventually accepted the male and spawning progressed. Females which had not ovulated, repeatedly rejected the male’s advances. The onset of spawning in the female was indicated by fluttering of the pectoral fins and successive horizontal undulations of the posterior trunk which served to sweep the spawning site. The laying of several thousand eggs was preceded by the female taking several large inhalations of water through the mouth and a slight raising of the posterior trunk. The male continued various behavior related to courtship while the female approached spawning. After spawning, the female smoothed the egg mass into a thin layer by using the posterior trunk and anal fin. After spawning, and while the female flattened the egg mass, the male copulated repeatedly. No copulation was observed prior to spawning. Semen was observed to be emitted from the tip of the penis and to leak from the female’s genital pore. Eggs were fertilized externally by the emitted sperm in the case of females which had no prior copulating experience in the present breeding season. The entire reproductive behavior of the pair ceased, as the female left the spawning site after flattening the egg mass. The male remained near by, but did not aerate the eggs. During the breeding season, the area around the female’s genital pore and along both sides near the base of the anal fin became slightly distended and soft by the accumulation of a serous fluid. It is presumed that this edematous modification serves to flatten the egg mass for softness without crushing the eggs.     

Females solicit sneakers to improve fertilization success in the bitterling fish (Rhodeus sericeus)

In most species there is greater variance in reproductive fitness among males than females. A consequence is the evolution of alternative male mating tactics, including sneaking behaviour whereby males attempt to mate with females courted by other males, which could undermine female choice. In bitterling (Rhodeus sericeus), fish that lay their eggs on the gills of living freshwater mussels, sneaking is common and successful. Here we show that female bitterling can significantly improve their fertility with an increase in the number of spawning partners. Females perform conspicuous behaviours associated with spawning more frequently close to sneakers, and spawn more eggs close to high-quality sneakers. This is the first time that females have been shown to engage in behaviour that increases the probability of sneaking, and raises the possibility of a sexual conflict over the number and source of ejaculates during spawning in this species.     

Females prefer mating males in the carmine triplefin,Axoclinus carminalis,a paternal brood-guarder

Synopsis This study investigates the role of male mating status in female choice patterns in the carmine triplefin, Axoclinus carminalis, a tripterygiid fish that exhibits paternal care. The distribution of daily reproductive activity is clumped, with many males receiving no mates and some receiving three or more. Females in this species do not prefer larger males, and characteristics of the oviposition site appear to have minimal effects on male mating success. When a female is removed from a male early in the daily spawning period, that male attracts fewer additional females for the remainder of the spawning period than does a control male. These changes in mating success are temporary, and do not affect mating success on subsequent days. A preference for mating males or males that are guarding eggs could provide asymmetric benefits for males to defend oviposition sites. This preference for males with eggs could be acting alone or with other factors such as high variance in oviposition site quality to favor the evolution of paternal care in fishes.     

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.     

Reproductive success in male sunbleak, a recent invasive fish species in the U.K.

The spawning behaviour of male nest guarding sunbleak Leucaspius delineatus , a recent invasive species in southern England, was studied and quantified for the first time. In the absence of physical differences between territorial and non‐territorial males ( i.e . colour, size, etc .), the reproductive behaviour of territorial males was analysed and related to reproductive success. The results showed that females preferred high‐courting and highly aggressive males. The initial cue in female mate choice, however, was based on courtship, while aggression was the decisive behavioural trait in influencing mate choice, providing a direct signal of physical condition and 'paternal competence'. Some males picked nest sites which were subsequently preferred by other males taking over the nest of a previous male ('communal nest'), with the new territorial male adopting the eggs already present at the nest. It appears that either due to female preference for nests already containing eggs or lower rates of sired egg predation by dilution among unrelated eggs, sunbleak males have adopted the mating strategy of allopaternal care.     

Female Mate Preference for Males Having Long and Symmetric Fins in the Bower-holding Cichlid Cyathopharynx furcifer

Female mate preference in a bower-holding cichlid, Cyathopharynx furcifer, was studied in Lake Tanganyika. Most males held territories with crater-shaped bowers in sand, but some males held territories without bowers. Territories were distributed adjacently and females visited them to spawn. After engaging in circling behaviour with the male, a female deposited eggs in the bower. Soon after spawning, the female picked the eggs up into her mouth and brooded them in places away from male territories. Female mate choice appeared to follow three steps: 1) females visited only bower-holding male territories, and more frequently visited territories of males that performed courtship displays at a higher frequency and had longer pelvic fins; 2) females preferred to start circling with males having longer and more symmetrical pelvic fins; 3) females chose males with more symmetrical pelvic fins as their mates. Less than 7% of females that visited male territories spawned eggs in the bowers. In contrast to other bower-holding species, bower size did not correlate with male reproductive success in C. furcifer. Bowers may therefore be essential as spawning sites or may function as a species recognition character for females. Female choice may be dependent instead on males having long and symmetrical pelvic fins apparent during the circling behaviour carried out in the bowers.     

Breeding pattern and nest guarding in Sicyopterus lagocephalus, a widespread amphidromous Gobiidae

Amphidromous gobies are usually nest spawners. Females lay a large number of small eggs under stones or onto plant stems, leaves or roots while males take care of the clutch until hatching. This study investigates the breeding pattern and paternal investment of Sicyopterus lagocephalus in a stream on Reunion Island. In February 2007 and January 2010, a total of 170 nests were found and the presence of a goby was recorded at 61 of them. The number of eggs in the nests ranged from 5,424 to 112,000 with an average number of 28,629. We showed that males accepted a single female spawning in the nest and cared for the eggs until hatching. The probability for a nest to be guarded increased with the number of eggs within it, suggesting that paternal investment depends on a trade-off between the reproductive value of the current reproduction and future nesting events. We showed that large nest stones were occupied by large males (TL >80 mm), whereas smaller males (TL <50 mm) were found under smaller cobbles, probably because of male–male competition for available nests. Our results suggest that the male’s choice relies upon a similarity to the female size, while the female’s choice was based on both body and nest stone sizes.     

Territoriality and reproductive behaviour in the goldsinny, Ctenolabrus rupestris L.

Territoriality and reproductive behaviour of the Goldsinny, Ctenolabrus rupestris, was studied in the waters around Tjärnö on the Swedish west coast. Observations were made by SCUBA-diving and from an observation raft and in aquaria.Patrolling, foraging, courtship and aggression were defined and quantified throughout the reproductive season and after. Spawning activity was also examined and it has been found that one male spawns with several females. During spawning accesory males may be present at fertilisation and their role is discussed. Both sinking and floating eggs were recorded, which are discussed.After spawning some females became territorial as well as some subadults. The territories showed a consistency according to their boundaries and occupants from year to year.     

条纹短攀鲈的交配习性和筑巢生境

条纹短攀鲈栖息于静止、植被茂密、靠近稻田和池塘边缘的浅水中。本文研究了它在泰国稻田和小池塘两种生境中的繁殖特征。与池塘相比,稻田筑巢地的pH值、溶解氧的含量和温度较高,而电导、溶解的固体物浓度较低,水也较浅。稻田中的种群密度低于池塘。雄性个体的体重和体长均大于雌性个体,且其体重与体长和体宽正相关,但雌性个体的体重仅与体长正相关;泡巢的面积与雄性个体的体长和重量均不相关。繁殖期间,条纹短攀鲈雌、雄个体交尾多次。每次交尾产卵3.74±1.02(Mean±SD)粒,每个繁殖季节交尾84±12.97次;在一个持续178±51.17min的产卵时段里,条纹短攀鲈平均产卵314±109.77粒。鱼苗孵化需30.75±0.55h,2日龄个体的大小平均为3.21±0.29mm。     

Observations on the breeding habits of Clarias gariepinus (Burchell)

Clarias gariepinus (Burchell) can be stimulated to spawn in aquaria by hormonal treatment and the breeding habits could be observed. After pre-spawning activities such as the raising of dorsal fins and chasing of the female, the male folds around the head of the female and small batches of eggs are laid, followed by some gas bubbles. Eggs are distributed by the female over a wide area. This behaviour agrees with field observations. The final stimulant to spawning under natural conditions is thought to be a substance called petrichor dissolved in flood waters that have run over dry ground.     

Spawning of the kissing loach (Leptobotia curta) is limited to periods following the formation of temporary waters

The kissing loach, an endangered species surviving only in a few Japanese rivers, spawns in the rice-field areas after migration from rivers in early June. To characterize the environmental conditions required for spawning of the kissing loach, spawning was assessed for two years both by direct observation of spawning behavior and by the appearance of eggs, larvae, and juveniles from June to October. All spawning of the kissing loach was limited to within a couple of days after the formation of temporary waters by remarkable rises in water level. Water temperature and daily rainfall fluctuated during the investigation periods, and no clear relationships with spawning were detected. Furthermore, all spawning was observed only in temporary waters with terrestrial grasses. Thus, spawning of the kissing loach is rigidly limited spatio-temporally to after the formation of temporary waters over terrestrial vegetation. Appropriate management of temporary waters will be crucial for the continued existence of this species.     

Synchronous female spawning and male mating behavior in a land-locked population of Japanese charr, Salvelinus leucomaenis japonicus

In resource-based promiscuous mating systems, synchronous spawning of females affects competition among males and variation in the reproductive success of males. We documented the mating behavior of Japanese charr (Salvelinus leucomaenis japonicus) through an annual breeding season to examine the relationship between female spawning synchrony and male mating behavior. Females spawned highly synchronously in the population studied, i.e., approximately half the spawning was finished within the first three days of the entire spawning season (11 days). The daily operational sex ratio (OSR) was nearly 1:1 through the spawning period. The number of males around a spawning female was very small (1.21+/-0.49 males per female) over the spawning ground and period, suggesting that a competitive male could effectively chase subordinate males away from a spawning female. A few males attempted to sneak near the oviposition site of females (16%; 9 of 57 breeding groups), while some males adopted sneaking tactics in the initial phase of females' spawning (24%). We did not observe any males to succeed in sneak fertilizations. We conclude that in this Japanese charr population, the synchronous spawning of females was related to the unbiased daily OSR, male aggregation around females, and consequently whether and how efficiently males engaged in sneak mating behavior.     

Shifting reproductive success in a shoal of Atlantic Cod, Gadus morhua L.

Individual and temporal variability in the spawning behaviour of Atlantic Cod (Gadus morhua L.) was examined by studying wild and reared individuals from the same putative population in a common spawning arena. The number of eggs fertilized per male was best explained by the number of aggressive interactions initiated by each male. A strong skew in male reproductive success and a temporal shift in male fertilization success were also observed. The latter co-occurred with an apparent similar temporal shift in the reproductive behaviour of the individual males. We hypothesize that energetic costs associated with reproduction, potentially mediated by sperm depletion in dominant individuals, may be responsible for temporal shifts in mating behaviour and reproductive success of male Atlantic Cod. We suggest that this hypothesis merits further study.     

The cost of peripheral males in a brook trout mating system

A focus on the reproductive contributions of males displaying alternative life histories has neglected the role of size-dependent peripheral males in salmonine mating systems. We documented mating behaviour of brook trout Salvelinus fontinalis, including observations of spawning, over two breeding seasons to determine the mating costs of peripheral males to dominant males (kleptogamy) and females (egg cannibalism). For males and females, the mating costs of peripheral males were substantial because more than half (56%) of all observed brook trout spawnings involved peripheral males. Males that paired with large females experienced a greater incidence of kleptogamy due to increased numbers of peripheral males present. Large males face a conflict when mating in that they prefer to spawn with large females; however, these same females attract numerous males against which the dominant male cannot defend. From paternity studies, we estimated that males that had peripheral males participate in spawning may fertilize, on average, equal numbers of eggs compared to males spawning solely with a smaller female. Females that paired with relatively smaller males had significantly more eggs eaten by peripheral males than females that paired with relatively larger males. Latency to spawn by females increased when paired with a relatively small male, and resulted in females obtaining a larger spawning partner. The observed patterns of size-assortative mating, kleptogamy and cannibalism are discussed in relation to mate choice for this population of brook trout. Copyright 1999 The Association for the Study of Animal Behaviour.     

Deceptive female oviposition behaviour elicits male ejaculation in the European bitterling

In European bitterling Rhodeus amarus , fish that lay their eggs in the gill chambers of living freshwater mussels, females perform conspicuous behaviours associated with spawning that increases the probability of males performing ejaculatory behaviour and participating in a spawning. A significant positive association was detected between behaviour in which a female performs a spawning action but without releasing eggs, here termed 'deceptive female oviposition', and ejaculatory behaviour by courting males.