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.     

Embryonic development and larval behavior of the kissing loach (<Emphasis Type="Italic">Parabotia curta</Emphasis>): adaptations to an ephemeral,hypoxic environment

Many teleost fishes in lowland fresh waters spawn in ephemeral flooded areas, the bottoms of which are prone to hypoxia. Little is known about how embryos and larvae deal with these potentially hostile environments. This study examines the functional and behavioral ontogeny of one such species, the kissing loach (Parabotia curta). Kissing loach eggs are demersal and adhesive. Hatching occurs at 24.8 ± 0.1 h post-fertilization at 25°C, much earlier than most fish species. The newly hatched larvae are precocious with no functional mouth, fins or eye pigmentation. Swimbladder inflation normally occurs at about 4 days posthatch, even before which the hatched larvae moved immediately toward the water surface to hang from water moss. Experiments with larvae 20 h after hatching showed that they spent significantly less time on the bottom in hypoxic water (2 mg/l) than in normoxic water, and suggest that hypoxia is a major directive factor in eliciting surfacing behavior. For the kissing loach, we have previously reported short-term spawning after the formation of flood areas as well as wide scattering of the spawned eggs in the temporal flooded areas. These traits with the present results of hatching at an early stage and the immediate upward movement of larvae are considered to be effective strategies for using ephemeral, hypoxic flooded areas for reproduction.     

Reproductive behavior of the territorial herbivoreStegastes nigricans (Pisces: Pomacentridae) in relation to colony formation

Reproductive behavior of the herbivorous damselfishStegastes nigricans was studied. Both males and females held individual feeding territories, which were distributed contiguously and formed colonies. Spawning season was from June to September. Females visited males' territories and spawned eggs on the algal nest. Males tended eggs until hatching but exhibited less care taking behavior than other damselfishes. The mating system was promiscuous in that both males and females spawn with several mates, even within a single day. However, only 1 female was accepted in the nest at a time. The restriction of the number of mates in the nest may be an adptation to prevent the sneaking which was often conducted by other males of the same colony, and may also be avoidance of disturbing the spawning female. Most spawnings were done among members of the same colony and inter-colonial spawnings were rare (27 cases of 333 spawnings observed). The results of this study suggest that colonial distribution ofS. nigricans may bring several benefits in reproduction to the colony members (e.g. easy mate findings within fellow colony members), but also seems to bring costs (e.g. restriction of number of mates).     

Reproductive and habitat isolation between two populations of the striated spined loach

Synopsis This study deals with ecological differences influencing reproductive isolation between two sympatric populations, the small and middle forms, of the striated spined loach. As natural hybrids are rare, isolation between these populations must be complete. There are some differences in their reproduction and early life history. Both reproduce in temporary waters flooded in summer, but their spawning sites are isolated from each other. The small form spawns in shallow areas of still water around irrigation creeks or fens. On the other hand, the middle form spawns in slow-flowing ditches. Spawning site segregation between them is an important isolating mechanism, and is a by-product of adaptation in their early life history.     

Simultaneous polygyny of the gobiid fish Asterropteryx semipunctata in relation to mate availability

Nesting males of Asterropteryx semipunctata conducted spawning behavior with 2–6 females simultaneously. We carried out field observations on a rocky reef in Kagoshima, Japan, to examine the hypotheses that large males will show multi-female spawning behavior because of their mating advantage, and that simultaneous multi-female spawning will occur when the operational sex ratio (OSR; the ratio of receptive males to receptive females) becomes female-biased. Contrary to our prediction, neither the total number of multi-female spawnings during a spawning season nor mean number of spawning females at a time were correlated with nesting male sizes. This indicates that larger males often did not conduct multi-female spawnings. As predicted, the incidence of multi-female spawning followed the change in the OSR over time—as the OSR in the study area became biased toward females, the incidence of multi-female spawnings gradually increased. Our results suggest that mate availability affects mating patterns in A. semipunctata.     

Effect of spawning number and ration on reproductive performance of the batch-spawning three-spined stickleback Gasterosteus aculeatus

An experiment quantified the effect of food ration and spawning number on the breeding season reproductive performance of batch-spawning, female three-spined sticklebacks Gasterosteus aculeatus . Individually housed females were maintained on fixed rations of 2, 4, 8 or 16% of their initial postspawning mass of live enchytraeid worms from immediately after their first spawning until they ceased spawning. Number of spawnings correlated positively with ration. Total breeding season reproductive investment (total wet mass of eggs produced as a percentage of initial postspawning female mass) ranged from 38% at the 2% ration to 147% at the 16% ration. At the lower rations, postspawning mass, batch fecundity and wet and dry masses of the batch declined over successive spawnings, with the rate of decline inversely related to ration. At the highest ration, there was no decline in batch fecundity and postspawning mass increased over successive spawnings. Mean reproductive investment per inter-spawning interval was higher at the highest ration, but at all rations declined over successive spawnings. Mean reproductive effort per inter-spawning interval (wet mass of eggs spawned as a percentage of the wet mass of food consumed over the inter-spawning interval) was inversely related to ration. At the higher rations, reproductive effort showed no trend over successive spawnings. Neither spawning number nor ration had a systematic effect on egg diameter, wet mass per egg, dry mass per egg or total lipid content of the eggs. If the rate of food intake was insufficient, although batch fecundity declined, the main adjustment was a reduction in the number of spawnings in the breeding season.     

The inter-spawning interval of the female Three-spined stickleback, Gasterosteus aculeatus

Female Three-spined sticklebacks spawn several times in a breeding season, at intervals of a few days. The changes that take place in females during the interval between successive spawnings were analysed. The average time between spawnings was 87.5 hours. Between spawnings females increased in weight by between 15 and 30%. The bulk of this weight increase was lost when the eggs were spawned. The increase in weight was because of an increase in weight of the ovaries, which showed an increase in both dry matter and water content. After spawning the ovaries formed 8 to 9% of the total body weight, and this increased to up to 24% just prior to the next spawning. The ovaries were maximally hydrated immediately after spawning. There were no significant changes in the calorific value of the ovaries, but the calorific value of the rest of the body was a minimum just after spawning. It was estimated that between spawnings females were making a net gain of energy at an average rate of 2 cal/hour. After spawning, courtship behaviour by the females waned, but recovered over the last half of the interval between spawnings. The inter-spawning interval of the Three-spined stickleback provides a useful situation in which physiological and nutritional aspects of reproduction in a teleost fish can be studied.     

Aspects of the spawning of labrid and scarid fishes (Pisces: Labroidei) at Enewetak Atoll,Marshall Islands with notes on other families

Synopsis Spawning of 32 species of Labridae and 13 species of Scaridae was seen at Enewetak Atoll, Marshall Islands. Most spawned on a reef bisecting the main ocean-lagoon passage which had strong tidal currents. Others spawned on lagoon reefs and in Halimeda beds. Polygynous haremic, lek-like and promiscuous mating systems were found which were species specific. Data on reproductive patterns, sexual dichromatism, sexual dimorphism, seasonality and spawning behavior were determined. Many spawned during the day in a time-phase dependent pattern from near sunrise to sunset. Scarid spawning began at slack high water or after when currents were starting to move out of the lagoon. Labrid spawning usually started about 30 min later with some continuing up to 2 h after high tide. With high tide before sunrise, scarid spawning began 30–50 min after sunrise as the current started flowing over the reef. With high tide near sunset, spawning occurred with an incoming current. Most labroids spawned on all phases of the moon. Acanthuridae (6 spp), Caesionidae (1 sp.) and Zanclidae (1 sp.) spawned after high tide at the same time as labroids. Pomacanthidae (5 spp.) spawned only shortly before sunset without reference to tidal currents. Fishes producing pelagic eggs at the lagoon-ocean channel spawned (1) at or slightly after high tide (44spp.), (2) in late afternoon without reference to tide (6 spp.) or (3) after slack low water (1 sp. ). Spawning style can vary within a single species in different environments. Despite the presence of many piscivores, no successful predation on spawning adults was seen. Predation on newly released eggs was uncommon. Labrichthys unilineata and Anampses twistii attempted to defend their eggs for a few sec after release. Attacks by piscivores on spawning adults on tropical reefs occur once per 100–1000 spawnings. Most are wary when preparing to spawn and prespawning behavior is easily interrupted. The risk from piscivores goes down and spawning ascent speed decreases with increasing size of spawners. Egg predation by zooplanktivores is less for pair spawners than group spawners possibly due to less conspicuous gamete clouds and times of spawning. Increasing height of egg release, speed and length of the spawning ascent, and trajectory alteration of ascending adults are believed to make it more difficult for zooplanktivores to locate eggs after release. For labrids, permanent full sexual dichromatism was found among haremic, lek-like and promiscuous mating systems. Species with temporary full dichromatism, permanent and temporary partial dichromatism and monochromatism were haremic. Smaller scarids were believed to have lek-like, and larger species haremic, mating systems. Smaller scarids had male looping behavior and post spawning displays, plus faster spawning ascents and different locations for egg release than larger ones. Eggs of 21 labrids were spherical or nearly spherical, ranging from 0.55–0.80 mm in diameter, and most had one oil globule. Among 7 is scarids, 6 had spindle-shaped eggs ranging from 1.25 ×0.50 mm 2.14× 0.48 mm while one had a nearly spherical egg. One scarid egg lacked an oil globule.     

Quantitative observations of predation during spawning rushes of the labrid fishThalassoma cupido at Miyake-jima,Japan

A rapid upward spawning rush is a nearly universal phenomenon among reef fishes that spawn pelagic eggs. Although spawning rushes have long been considered to have evolved as a defense against egg predators and/or attacks on the spawning fishes by piscivores, these hypotheses have never been tested in the field. We analysed piscivore attacks during three motor patterns associated with group spawning of the labrid fishThalassoma cupido at Miyake-jima, Izu Islands, Japan. Egg predation on group spawnings was also quantified. Of 206 piscivore attacks on spawning fishes by seven predator species, 17.1 (83.1%) occurred during 461 spawning rushes (1/2.7 spawning rushes). No attacks were successful during spawning rushes, and only four kills were made in 206 attacks summed from all three motor patterns associated with spawning, amounting to a piscivore success rate of only 1.9 %. In contrast, gametes from 90 of 213 spawnings (42.3 %) were consumed by nine species of planktivorous fishes. Spawning fish seemed not to recognize egg predators and made no attempt to avoid them, often spawning in the midst of waiting aggregations of damselfishes. Our data indicate that the rapidly of the spawning rush ofT. cupido serves as an excellent defense against piscivores, but is ineffective against egg predators. Evidence is presented from observations of seven other species suggesting that spawning ascents provide little protection against water column egg predators in predator-rich environments.     

Spawning habitat in the spined loach, <Emphasis Type="Italic">Cobitis taenia</Emphasis> (Cypriniformes: Cobitidae)

Spawning habitat of the endangered spined loach Cobitis taenia was studied in northwestern Germany. The distribution of eggs of the loach in the field was best correlated with the density of vegetation but had little correlation with current velocity, water depth, or bottom substrate. In the aquarium, the loach chose dense vegetation for an oviposition site, as inferred by the direct observations of spawning and the location of spawned eggs. It is concluded that such habitat specialization is an important element in the autecology of the endangered fish and a critical stage in the habitat requirements of its populations.     

Scatter-spawning of the catfish,Silurus asotus

Reproductive behaviour of the catfish,Silurus asotus was studied in temporary waters around paddy fields. Spawning occurred nocturnally during the first week from the initiation of irrigation. In reproductive activities, a male first energetically pursued a female with its head near to the female’s belly (chasing) and then began to cling to the female’s body from the side, bending its tail or head (clinging). Finally the male enfolded the female’s body, with its anus near to the female’s (enfolding). In some cases, 2–4 males pursued a single female and two males enfolded a female at the same time. Although no aggressive behaviour was evident between males, it was always the largest male that could most frequently approach and enfold the female. The mating pair moved a long distance in a ditch, paddy field and/or creek, performing reproductive activities. It is thought that the spawning site and period of spawning of the fish enable the larvae to avoid the danger of predation and to efficiently feed, firstly on plankton and later on larvae of other fishes which become abundant during the irrigation period. Although some eggs and larvae may die due to the drying out or high water temperatures of such unstable temporary waters, scattering eggs may reduce the incidence of the annihilation of the young.     

Spawning behavior and reproductive isolation of two species ofPseudolabrus

The reproductive behavior of two wrasses,Pseudolabrus eoethinus (Richardson, 1846) andP. sieboldi Mabuchi & Nakabo, 1997, was studied on a rocky coast in Shikoku, Japan, where the two species occur sympatrically. In the study area, individual males of each species established mating territories within which pair spawnings occurred. Pair spawnings were observed only between conspecifics, indicating that the two coexisting species were reproductively isolated. Both species usually spawned during the afternoon in early winter and had similar reproductive behavior. Although habitat, temporal and behavioral isolation mechanisms were not apparent, reproductive isolation seemed to occur via premating isolation through visual recognition of conspecific mates because the two species differed in body size and coloration. On one occasion a maleP. sieboldi was observed to perform streaking behavior to join a spawning pair ofP. eoethinus. Thus, postmating isolation between the two species is also discussed.     

Reproductive ecology of the Far Eastern catfish, Silurus asotus (Siluridae), with a comparison to its two congeners in Lake Biwa, Japan

Spawning activity of the catfish, Silurus asotus, takes place in temporary water (rice fields) from early evening to midnight, generally in connection with rainfall, from late April to late August. Spawning of the catfish was correlated with hydrographic parameters dependent on rainfall: daily precipitation, turbidity, water depth, and water temperature. The spawning habits of the catfish, and in particular the use of temporary waters such as rice fields that become submerged after rainfall or by irrigation, are presumed to be adaptations to the Asian monsoon climate with a pronounced rainy season. The apparent sex ratio of the catfish was extremely biased toward females. Intraspecific variation in the reproductive ecology, particularly mating behavior, of this species is observed among local populations. Factors that may have caused this variation are discussed in the context of a comparison of mating behavior, reproductive environment, and sex ratio between the Lake Biwa population of S. asotus and other conspecific populations, as well as two other species of silurid catfish that occur in the Lake Biwa drainage, S. biwaensis and S. lithophilus.     

Male reproductive competition in spawning aggregations of cod (Gadus morhua, L.)

Reproductive competition may lead to a large skew in reproductive success among individuals. Very few studies have analysed the paternity contribution of individual males in spawning aggregations of fish species with huge census population sizes. We quantified the variance in male reproductive success in spawning aggregations of cod under experimental conditions over an entire spawning season. Male reproductive success was estimated by microsatellite-based parentage analysis of offspring produced in six separate groups of spawning cod. In total, 1340 offspring and 102 spawnings distributed across a spawning season were analysed. Our results show that multiple males contributed sperm to most spawnings but that paternity frequencies were highly skewed among males, with larger males on average siring higher proportions of offspring. It was further indicated that male reproductive success was dependent on the magnitude of the size difference between a female and a male. We discuss our results in relation to the cod mating system. Finally, we suggest that the highly skewed distribution of paternity success observed in cod may be a factor contributing to the low effective population size/census population size ratios observed in many marine organisms.     

Lunar synchronization of spawning in sea bass, Lates calcarifer (Bloch): effect of luteinizing hormone-releasing hormone analogue (LHRHa) treatment

Based on egg collection records, spontaneous spawning activity of sea bass, Lates calcarifer (Bloch), reared in floating net cages followed a semilunar cycle. The peak of multiple spawnings coincided with declining spring tides of quarter moon periods. Maximum diameter of intra-ovarian, ripe oocytes (0.51–0.55 mm) occurred in synchrony with the quarter moon periods. Smaller oocytes (0.44–0.47 mm) were sampled during the new and full moon periods. Two structural analogues of luteinizing hormone-releasing hormone (LHRHa) (des-Gly¹⁰, d -Ala⁶-LHRH ethylamide and d -Ala⁶, Pro⁹-Nethylamide-LHRH), in pelleted or dissolved form, induced mature female sea bass with a mean egg diameter of at least 0.40 mm to spawn at any day during the lunar cycle. The onset of spontaneous and LHRHa-induced spawnings occurred during low tides in the evening until dawn (from 19.00 to 05.00 hours). These results demonstrate that LHRHa can effectively induce mature sea bass to spawn independent of the highly predictable semilunar spawning rhythm. In addition, the occurrence of both spontaneous and hormone-induced spawnings at a precise period of the day suggest a tidal and diurnal cue entraining spawning of mature female sea bass.     

Age, growth and reproduction of Cobitis paludica in a seasonal stream

In the Iberian endemic loach Cobitis paludica , in a small seasonal tributary of the Guadalquivir River, 65–70% of the total growth in length occurred in the first year of life. The maximum ages observed were 3+ years in males and 4+ years in females. Both sexes matured at the beginning of their second year of life. There was a significant difference from 1: 1 in the overall sex ratio of 412 males to 674 females. Spawning began in late March and ended in July. Cobitis paludica is a multiple spawner that releases a minimum of two batches of eggs per female each year. Once spawning had started, however, there was no recruitment from the stock of transparent oocytes to the mature stock. The number of eggs decreased from the first batch to the second, but there was no significant difference in the mean egg diameter between batches. Compared with other populations of this species, the present population, from a low latitude, is characterized by a low number of age groups, fast growth, early maturity and high fecundity in multiple spawnings. These life-history characteristics are typical of species in unstable environments, where adult mortality is high, variable or unpredictable.     

Predictable and unpredictable spawning events: in situ behavioural data from free-spawning coral reef invertebrates

Summary

We describe the spawning behaviour and some aspects of spawning periodicity in a diverse group of marine invertebrates, principally echinoderms, but including sponges, anthozoans, molluscs, and polychaetes. Our observations were made both opportunistically and on a systematic basis between 1978 and 1992 on the central and northern Great Barrier Reef. Spawning was predictable in some of the species observed, for example Bohadschia argus, Euapta godeffroyi, and Stichopus variegatus (Holothuroidea), which exhibited regular lunar and diel periodicity. Others, such as Holothuria coluber, Actinopyga lecanora, and Bohadschia graffei (Holothuroidea), Acanthaster planei (Asteroidea), Hyotissa hyotis and Arca spp. (Bivalvia) exhibited no clear lunar or diel periodicity in spawning behaviour. Mass heterospecific spawnings which involved several species, often from different phyla, were commonly observed. The species participating were usually those with unpredictable spawning patterns and while the species involved were diverse there were also occasions when spawning involved species from the same genera. Fertilization rates were measured in situ for the predictable spawner Bohadschia argus and were found to vary between 0–96% depending on the circumstances of the spawnings. Fertilization rates for the unpredictable spawners showed similar variability; Holothuria coluber and Actinopyga lecanora ranged from 9–83%.     

Reproductive behavior and mate fidelity in the monogamous goby,Valenciennea longipinnis

Reproductive behavior and mate fidelity of the gobiid fish,Valenciennea longipinnis, were studied on the coral reef at Sesoko Island, Okinawa, Japan. These fish usually live in pairs, not only foraging together for benthic animals in sandy areas, but also constructing several burrows within their home range. Before spawning, both fish, although mainly the male, constructed a mound, piling up dead-coral fragments, pebbles, shells, sand and algae onto one of the burrows. After spawning an egg mass on the ceiling of the burrow, the female stayed outside and continued the construction and maintenance of the mound for 3–5 days until hatching, while the male tended the eggs inside. Mate guarding of females seemed to prevent males from monopolizing several females. Although some pairs showed mate fidelity through several spawnings, more than half of the pairs broke up after only one spawning. The pair bond was broken by mate desertion and the disappearance of each sex. Both sexes preferred larger spawning partners; larger females spawned more eggs and larger males provided better egg care. Mate desertion occurred when larger potential mates, relative to the current partner, became available. The frequency of solitary individuals was higher in males than in females, resulting in females deserting their mates more often than males. Two factors seem to have facilitated mate desertion: (1) occurrence of size mis-matched pairing and (2) overlapping home ranges.     

Spawning habits of the fluvial eight-barbel loach Lefua sp. observed in captivity

The spawning behavior and reproductive characteristics of the fluvial eight-barbel loach (nagare-hotoke-dojo) Lefua sp. (sensu Hosoya, 1993) were examined in 11 male–female pairs of 31.5–61.5 mm in standard length (SL) (males) and 34.1–70.9 mm SL (females) in experimental tanks, each with a layer of charcoal sand. Spawning of Lefua sp. began at a water temperature between 13.4° and 15.9°C. For each pair, on the days when spawning occurred, the male was observed to patrol the bottom briskly, whereas the female swam up and down along the tank glass. Just before spawning, the females attempted to burrow into the sand or under a stone using beats of the caudal fin. When stimulated by being pecked at or being bitten by the male, the female redoubled her effort to find cover and the male followed her, after which they finally succeeded in finding cover. Muscular spasms, resembling trembling, were observed and they remained adjacent to each other. A moment later they both swam up out of the sand, completing the spawning. Each spawning event with a sequence of these behaviors took place within a 24-h cycle, with mating behavior not being observed to continue into a second day in this study. A total of 69 batches of eggs laid were confirmed in the experiments. Lefua sp. showed basically batch-spawning to a maximum of 13 batches. With the increasing number of batches, the number of eggs spawned in each ensuing batch decreased and the interval between spawning events lengthened. The spawned eggs tended to increase in diameter as the size of the female increased.     

Spawning frequency, fecundity, egg weight and spawning type of silver pomfret, Pampus argenteus (Euphrasen) (Stromateidae), in Kuwait waters

Silver pomfret, Pampus argenteus, were collected by fishing with drift gillnets on one spawning ground in Kuwait waters during 1998–2000. Fish size frequency, sex ratio, maturation cycle, spawning frequency, fecundity and egg weight were assessed. The length–weight relationship differed between sexes whereby females were significantly bigger than males. Spawning started in mid‐May and continued until early October. During this time the water temperature ranged from 26.0 to 32.8°C, salinity was ? 39.0‰ and water depth ranged between 5 and 12 m. Large females spawned earlier than young spawners and the overall percentage of males during the spawning period was 70.3%. Spawning occurred after 13.00 h, with peak spawning between 15.00 and 18.00 hours during outgoing tide. Mean daily spawning frequency amounted to 63.2%. Spawning activity was found to be associated with the lunar cycle and spawnings were concentrated during the first and third quarters of the moon period, indicating a semilunar reproduction cycle. It was concluded that a female would spawn at least six times during the season. No change was observed in relative fecundity during the peak spawning season (June–August). Average relative batch fecundity was 176.3 eggs g^?1 somatic weight (SW), corresponding to a relative total fecundity of 1058 eggs g^?1 SW, which is 1.5 times higher than estimates obtained from counting the standing stock of oocytes. Bigger fish produced heavier eggs and the egg weight decreased as the spawning season progressed. Based on gonadal cycles, oocyte size frequency distribution and total fecundity, we concluded that silver pomfret is a multiple batch spawner with indeterminate fecundity.